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Gardens’ Bulletin 


Singapore 


~ VOL. 62 (1) 2010 ; ISSN 0374-7859 


THE GARDENS’ BULLETIN SINGAPORE 


The Gardens’ Bulletin Singapore publishes original papers on plant taxonomy 
(including revisions), horticulture, phytogeography, floristics, morphology, 
anatomy and related fields with emphasis on plants in the West Malesian 
region. 


. Dr. B.C. Tan Dr. Jana Leong-Skorni¢kova 
Singapore Botanic Gardens Singapore Botanic Gardens 
(Editor) (Assistant Editor) 
De PY. Tan Ms. C. Soh 
National Parks Board Singapore Botanic Gardens 
(Assistant Editor) (Journal Business Manager) 
EDITORIAL BOARD 

Dr. S.C. Chin Dr. M.C. Roos 

Singapore Botanic Gardens National Herbarium Netherlands 
Singapore Leiden University, The Netherlands 
Dr. M.J.E. Coode Dr. E. Soepadmo 

Royal Botanic Gardens Forest Research Institute Malaysia 
Kew, U.K. Kepong, Malaysia 

Prof. Sir P. Crane Prof. T. Stuessy 

Yale University University of Vienna 

New Haven, USA Austria 

Dr. R.T. Corlett Dr. W.K. Tan 

DBS, National University of Singapore National Parks Board 

Singapore Singapore 

Dr. W.J. Kress Dr. I.M. Turner 

Department of Botany, NMNH Research Associate 

Smithsonian Institution Singapore Botanic Gardens 
Washington DC, USA Singapore 


The Gardens’ Bulletin is published twice yearly by the National Parks Board, Singapore. 
Neither the National Parks Board nor the Editorial Board is responsible for the opinions or 
conclusions expressed by the contributing authors. 


The annual subscription for the Gardens’ Bulletin is Singapore $100.00 including 
postage. Overseas subscribers are required to make payment in the form of bank drafts 
or international money orders in Singapore currency payable to National Parks Board, 
Singapore. 


Instructions for contributing authors are found on the inside back cover. 


[Cover photo: Inflorescence of Curcuma vitellina (see p. 114); photo by Jana 
Leong-Skorni¢kova. ] 


VOL. 62 (1) 2010 


ot DEC 21 2010 


The Gardens' Bulletin 


Singapore 


CONTENTS 


Ahmad Sofiman bin O. and P.C. Boyce 
Studies on Monstereae (Araceae) of Peninsular Malaysia I: Rhaphidophora 


(MEV ALIN ALAAMO WIPE ECOL OiOR WIESE NI Al aY SIA sores scrapers eace ses u- si ce-wancsessassiesuaserscescsecsestccesetes 


Ahmad Sofiman bin O., P.C. Boyce and Chan L.-K. 
Studies on Monstereae (Araceae) of Peninsular Malaysia III: Scindapsus lucens, 


a New Record for Malaysia, and a Key to Peninsular Malaysian Scindapsus..............:.00+- 


Ardi, W.H. and M. Hughes 
Begonia droopiae Ardi (Begoniaceae), a New Species of Begonia 


RE GUELTR VES ER ULET cl Rites eee ce cc cece eee eee Sosa sco suceo lececawesueny sos vavabeunendocabossascecsaatevees 


Arifiani, D. 
Newly Recorded Endiandra R. Br. (Lauraceae) from Waigeo Island, Raja Ampat, 


AGU AMMUTT CHOTA Scene esas eee ee cee etna evcn ene veseeccnuscccanncsodesu cane sasstesteccacereaessseseee 


Atik R. 
Species of Marasmius (Agaricales: Tricholomataceae) from Kayan Mentarang 


IN AEC IT Ale ya Konya CATT ATI GANS Gl OMESTA oe ce cccsescacsoceovwovsacnvecasacdecscesarcecesesessscosencesscenccteees: 


Hadiah, J.T. and B.J. Conn 


Lectotypification of Elatostema subscabrum H.Schroet. (Urticaceae) .........:eeeeeeeeee 


Haerida, I., S.R. Gradstein and S.S. Tjitrosoedirdjo 


Lejeuneaceae subfamily Ptychanthoideae (Hepaticae) in West Java «0.0... 


Kurzweil, H., S. Watthana and S. Lwin 


Phaius takeoi (Orchidaceae) Newly Recorded from Thailand and Myanmat ................... 


Leong-Skornickova, J., Tran H.-D. and M.F. Newman 


Curcuma vitellina (Zingiberaceae), a New Species from Vietnam ..........:.:ccceeseeteeeteeeeeees 


Lindsay, S. 


New Combinations in Haplopteris (Adiantaceae) for the Flora of Peninsular Malaysia... 


Nguyen, V.D., J.C. Regalado, Jr. and Vu T.C. 


A New Species of Alocasia (Araceae-Colocasineae) from Cambodia ............:seeeee 


ARNOLD A2B0RER ae] 
UBRARY 


ISSN 0374-7859 


ore 9 


43 


119 


121 


Nor-Ezzawanis, A.T. 
New Combinations VMalaysianis tap biylea cea ce sereeneeete teers cane nen veserse te seated seer et enree sans 127 


Poulsen, A.D. and C.D. Specht 
JN INDY S| VSOES Olt (COMES 1H KOA BOTAN EO) ecccecencaseospeccecnccerconsecontcnos acc -peccoconaaconnSotcenocecpenanancec 135 


Rajbhandary, S.,M. Hughes and K.K. Shrestha 
Three New Species of Begonia Sect. Platycentrum from Nepal ..............:csscccssesesesseseseneeees 143 


Sudarmono and B.J. Conn 
Genetic Variation of Populations of Scutellaria slametensis and S. discolor 


(Lamiaceae) on Gunung Slamet, Jawa Tengah (Indonesia) ............::::ceeccescsssseeseseeseeecseesenees 155 
Turner, I.M. 

A new species of Polyalthia (Annonaceae) from Sabal «.............200.ccsccssscsececcesseeseesseossosseesece 173 
Wong S.Y. 

Studies on Schismatoglottideae (Araceae) of Borneo XIII: A Revision of the 

SGI TRY UCT ONRMOPMAKY HA MUON! SI OASIS (COO) OES. cc rooecence-eeceecoccoccoreracecoccececoccocchLce ce pococtaASHocenSDOE SOS 177 


Book Review 
Flora of Peninsular Malaysia, Series I. Ferns and Lycophytes. Vol. 1. by B.S. Parris, R. Kiew, 
R.C.K. Chung, L.G. Saw & E. Soepadmo (eds). 
TE 1 Pal 24 (Opa 7 DEON BREET Oto dceoancoosoDODSOOOODNs0NS 2A 


Date of publication: August 2010 


Published and copyrighted by 


National Parks Board 
Singapore Botanic Gardens 
1 Cluny Road 
Singapore 259569 


Printed by Oxford Graphic Printers Pte Ltd 


Gardens’ Bulletin Singapore 62 (1): 1-8. 2010 l 


Studies on Monstereae (Araceae) of Peninsular Malaysia II: 
Rhaphidophora latevaginata, Newly Recorded for 
West Malaysia 


AHMAD SOFIMAN BIN OTHMAN AND PETER C. BOYCE * 


Pusat Pengajian Sains Kajihayat 
Universiti Sains Malaysia 
11800 USM, Pulau Pinang, Malaysia 
*Corresponding author: phymatarum@gmail.com 


Abstract 


Rhaphidophora latevaginata M.Hotta, a neotenic, shingling, climbing aroid, 
hitherto considered a Bornean endemic, has recently been found and 
collected from the southern part of the east coast of Peninsular Malaysia 
(Johor: Kota Tinggi and Mersing), where so far it appears to be restricted 
to kerapah and the drier (raised podzol) facies of seasonally inundated 
peatswamp forest. This discovery of R. /atevaginata takes to 18 the number 
of Rhaphidophora known to occur in Peninsular Malaysia, of which three 
are endemic. An updated description of R. latevaginata, a key to the 
Rhaphidophora species of Peninsular Malaysia, and a plate illustrating the 
diagnostic characters of those with shingle-stage juveniles is presented. A 
brief note on the significance of the new record with regard the Riau Pocket 
is made. 


Introduction 


Since the publication of an alpha-taxonomy for Peninsular Malaysia (Boyce, 
1999). and the Peninsular-relevant taxonomic alterations made for Borneo 
(Boyce, 2001), further study of Rhaphidophora in Peninsular Malaysia 
has generated additional data that sheds light on possible biogeographical 
patterns not earlier apparent. Most recently the discovery in Perak of R. 
megasperma Engl. (Baharuddin & Boyce, in press), previously regarded as 
a Bornean endemic and furthermore belonging to a species group until now 
considered to be restricted to E Sunda, Papuasia, and the tropical Western 
Pacific, and now the discovery of R. latevaginata in Johor, is providing 
compelling non-woody plant support to the Riau Pocket phytochore (Corner, 
1960; Ashton, 2005). All terminology used here follows Boyce (1999). 


— — 


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Gard. Bull. Singapore 62 (1) 2010 


Key to Rhaphidophora in Peninsular Malaysia (adult plants) 


. Leaf lamina variously pinnately divided and/or perforated ...............:esceeseeseeeeeees 2 
- Leaf lamina Cmte. is: si.i2..ccsecducascvesantie tessoee oe tae te oansateol cos canes: evs tee eudaegussaceacnde costeemn ere 9 
Leaf lamina abaxially pubescent, especially the mid-rib and primary lateral veins.....3 


Leaf lamina abaxially pla brotis:c.22:.jcccc. cede cccxecsee cece oe ceceseere ace ee 4 


. Plants flowering on adherent stems; mature leaves with numerous perforations 


along both) sides of the mtd nib x2. 22. -sssescssncee nen - sesecee a saseeeee ee eee eee R. foraminifera 


. Plants flowering on free stems; mature leaves with one to three perforations on 


one side of the mii rilb) ia. cs ccscesesoce-neesten as tt cps cece etna ae eee cee R. puberula 


Rheophytes; leaves of flowering plants occasionally entire.................... R. beccarii 


. Medium to high-climbing lianes; leaves of flowering plants never entire.............. 5) 


. Active shoot apices with sparse to copious netted fibre; feeding roots conspicuously 


ramentose-scaly; lamina of mature plants pinnatisect, the pinnae often perforated 
basally and appearing stilted. Juvenile plants with leaves overlapping in the manner 
ofroor shingles (Shingle: ClimDErs) he ccc. csee tte cee eee R. korthalsii 


. Active shoot apices glabrous; feeding roots smooth or slightly corky-lenticillate; 


lamina of mature plants variously split and perforated but never with the pinnae 
perforated basally and appearing stilted. Juvenile plants not shingle climbers)...6 


Leal lamina.at leastpartiallly pom ates. -c-ocerecseeecceee aces eee ee 6 
. Leaf lamina not pinnate, either entire or with a few, large ovate to rhomboid or 


trapezoid perforations, these sometimes extending almost to the margin............. 8 


.Leaf lamina up to 53 x 105 cm; sparsely to rarely + entirely pinnatipartite,or 


pinnatisect; petiole 40-70 cm long, petiolar sheath extending % - %4 along petiole; 
spadix up to 14 x 2 cm, stoutly cylindrical, inserted decurrently from 2 cm on peduncle; 
plants:exclusively of montane forest: se ccece tenes eeeee eee R. nicolsonii 
Leaf lamina up to 42 x 38 cm but usually much smaller, sparsely to + entirely deeply 
pinnatipartite to nearly pinnatisect, occasionally with large rhombic perforations 
adjacent to mid-rib; petiole 10-34cm long, petiolar sheath extending to base of apical 
pulvinus, inflorescences a few together, spadix up to 3.5 x 1 cm, cylindrical, inserted 
slightly decurrently on peduncle; plants of lowland forest.............. R. tetrasperma 


. Inflorescence two (sometimes more?) together, each subtended by a prominent, 


soon falling, cataphyll, and arising from an elongated reiterative floral sympodium 
at the tip of a plagiotropic free lateral shoot; spathe caducous; stigma impressed 
irregularly elliptic, longitudinally ontentated in cc... cree ere eee R. corneri 


. Inflorescence three or more together, each subtended by a prominent persistent, 


later marcescent cataphyll, and arising from an much abbreviated reiterative floral 
sympodium arising laterally on a climbing shoots; spathe marcescent; stigma raised, 
PUT CHLORINE ones. 50s dcdsancaenscoanere <oseueseeevec hes oone Seem eens oe eee eo R. megasperma 


Studies on Monstereae (Araceae) of Peninsular Malaysia II 3 


9. Leaves always shingling, even in flowering individuals; leaf laminas stiffly coriaceous, 
broadly oblong-ovate-elliptic, 848 x 6.5-20.5 cm, bright green, slightly to markedly 
glaucous, base truncate-cordate to broadly cuneate. Flowering on clinging shoots 
Pe ee rane eee aac ac ey connie reusing ssundesesdes sdesuoesnsseassssieeshosuseessacakdouews R. latevaginata 

9. Leaves spreading in adult and flowering individuals; leaf laminas variously coloured 


but never glaucous. Flowering on free or clinging ShHOOTS.............c:ecceseeeteeteeees 10 
10. Abaxial surface of lamina and apical pulvinus pubescent.................. R. puberula 
Mabamind anGapical PUlvinUs MOL AS-ADOVE...z.......<....c--censecessnesacsnssssesnnerteseonsedsatsacs 11 


11. Flowering shoots consisting of scattered fans of large litter-trapping leaves carried 
on short stout shoots and held at about 90” to the + leafless main stem................ 
ae SE tee Ae Sea oe cans cn satvandeqhdsnecsdustaidesnoenpcaestseateaveaueoeadeswedecetens R. angustata 


GND OMe GIMPISMOOUS MOLI AS AD OW Cxco2.2520.520:.0.acdsascnseuccessvscosueccveccnsseecsecstetsaseoesbeedaesonced oz 12 
12. Plant climbing; feeding roots smooth, or minutely asperate........... eee 13 
12. Plant rheophytic; feeding roots conspicuously ramentaceous............... R. beccarii 


13. Stems sub-terete to weakly 4-angled, scabrid to asperous, older portions with thin, 


brittle pale brown epidermis; spathe exterior minutely puberulent......... R. lobbii 
13. Stems conspicuously 4-angled, smooth, older portions remaining green; spathe 
EE NAMEN AS EONS Bes eo os ns on Ses nas atnsabavoeisAuve susedibaceeoanseeswaveeteseasvoseacecsesvinveats 14 


14. Apices of active stems with netted prophyll, cataphyll and petiolar sheath remains 
SAS TASC a severe rece a csc SSST SELL cus SSoasee ve evensoscecesusdvessessisvsonsacavssvessueecieessies R. maingayi 


Pes MAIC OMAC (IVC SU CMS TI AKC Glew sias2covnsn sien ieecenseanccoeceanrsnest-cosecsccsncsovsscuecoessssetsesedseceee 15 
iSaspadix at anthesis 9-20 cm long, tapering apicallly.....................:cce-ceoresesseeseosonees 16 
15. Spadix at anthesis 3.5-6 cm long, cylindrical to weakly clavate...........c:cceeee 17 
igropadin 9-iticna lone; Samdstonme Se PTAMILE............2.200csseissseooesesccoseeneeeee R. montana 
16s; Spadix 15-20 cm long; limestone obligate......:........:.s:.s-csssessessecessens R. burkilliana 
1s SS ORYC TDS (57 SAG Ui Oe eee er Ee a PS Pe 18 
RNiemS a AG aI ANY CAKINICIAN AL Coes ake canc eaters cessavececeaneecvtvevesertorsoectsbeenecteuseosessecevnenes R. sylvestris 


18. Leaf lamina thickly coriaceous to almost fleshy, falcate-elliptic- lanceolate to 
falcate- oblong or falcate-oblanceolate, 4.5-25.5 x 1.5-5 cm; margins slightly reflexed, 
this becoming greatly accentuated in dried material.................0004 R. crassifolia 

18. Leaf lamina thinly coriaceous, narrowly falcate-elliptic to falcate-lanceolate 
or falcate-oblanceolate, 2.5-16 x 1.2-3 cm, margins flat, leaf drying pale straw- 
CONIA sees ca ca wpe ence sab os soesacU vas osnstnap ac cosceseedsscdsnssbbeesdéadpesaasatnconaeesas R. minor 


4 Gard. Bull. Singapore 62 (1) 2010 


a 


Plate 1. A-B. Rhaphidophora latevaginata M.Hotta. A. Leaf (abaxial view) showing the 
strongly unequal wings of the long-persistent petiolar sheath. Note that the sheath is mainly 
adnate to the moderately short petiole, with only a short free-auriculate portion. B. Juvenile 
plant. C-D. Rhaphidophora kothalsii Schott. C. Leaf (abaxial view) showing the equally winds 
to the swiftly-marcescent petiolar sheath. Note that the sheath is mostly free from the very 
short petiole, with the greater part forming a long, narrowly triangular, free-ligulate portion. 
D. Feeding root showing the diagnostic ramenta. Images © P.C.Boyce 


Studies on Monstereae (Araceae) of Peninsular Malaysia II 5 


Rhaphidophora latevaginata M. Hotta, Acta Phytotax. Geobot. 22: 44 
(1966); Boyce, Gardens’ Bulletin Singapore 53: 51-54, Fig. 10 (2001). -Type: 
Malaysia, Sarawak, Bintulu (‘4th Division’), Bintulu District, about 4 km 
east from Minah Camp, Sg. Kakus, 4 Oct 1963, Hirano & Hotta 140 (KYO, 
holo!). Plate 1A & B. 


Moderate to very large, robust, pachycaul, homeophyllous neotenic liane to 
12 m; seedling stage a non-skototropic shingling juvenile shoot; pre-adult 
plants forming small terrestrial colonies of shingling closely appressed 
leaves; adult shoot architecture comprised of clinging, physiognomically 
unbranched, shingling to very densely leafy, sterile stems and almost identical 
fertile stems. Stems weakly compressed-terete to weakly rectangular in cross- 
section, smooth, pale green, without prophyll and cataphyll fibre, internodes 
to 12 x 2 cm, separated by prominent straight scars, but scars obscured by 
leaf bases on all but the oldest stems, lower parts of stem later sub-woody 
with slightly shiny cracking thin pale brown epidermis; flagellate foraging 
stems moderately well developed although often somewhat short and leafy; 
clasping roots arising densely from the nodes and internodes, prominently 
scaly; feeding roots ca 3 mm diam., brown, minutely pubescent. Leaves 
distichous, appressed, ascending and shingling, becoming slightly scattered 
and spreading (often litter-trapping) towards fertile tips; cataphylls and 
prophylls membranous,soon drying black and persisting briefly before falling; 
petiole deeply canaliculate and winged, 3-22 x 0.5-2 cm, smooth, apical and 
basal pulvinus obscure in young leaves, later becoming prominent, especially 
the basal pulvinus; petiolar sheath very pronounced, up to 2.5 cm wide, 
rather thickly membranous, adnate to the petiole for much of their length, 
prominently rounded short-auriculate, especially the larger (outer) sheath, 
outer sheath (away from climbing surface) greatly expanded and partly to 
completely obscuring stem, both sheaths persisting some considerable time, 
much later (and then almost exclusively on adult plants) rotting to produce 
two large scars extending to the top of the petiole; lamina broadly oblong- 
ovate-elliptic, 8-48 x 6.5-20.5 cm stiffly coriaceous, bright green, slightly to 
markedly glaucous, base truncate-cordate to broadly cuneate, very briefly 
decurrent, apex rounded to acute with a tiny apicule; midrib prominently 
raised abaxially, slightly sunken adaxially; primary venation pinnate, slightly 
raised abaxially, more so adaxially; interprimaries sub-parallel to primaries, 
slightly raised on both leaf surfaces; secondary venation tessellate-reticulate, 
slightly raised abaxially, + flush adaxially, all veins much more prominent in 
dried material. Inflorescence solitary on a clinging shoot, subtended by a 
fully developed foliage leaf and one or more cataphylls; peduncle laterally 
compressed-cylindrical, 6.5-11 x 0.5-0.7 cm; spathe not observed; spadix 
stoutly cigar-shaped, sessile, inserted + level on stipe, 17.5 x 1.5 cm, pale 


6 Gard. Bull. Singapore 62 (1) 2010 


green; stylar region rhombohexagonal, ca 2 x 1 mm, truncate; stigma slightly 
raised, elongated, longitudinally orientated, ca 0.75 x 0.2 mm; anthers not 
exserted at male anthesis; infructescence stoutly cigar-shaped, 15 x 2 cm, 
stylar region becoming convex at fruit maturity. 


Specimens seen: MALAYSIA. Johor Bahru, Mersing, Kluang - Mersin Road, 
km 39, 02° 15’ 78.2”; 103° 43’ 79.2” 56, 18 Apr 2010, PC Boyce, Siti Nurfazila 
Abdul Rahman & Ooi Im Hin AR- 3039 (KEP); Johor Bahru, Kota Tinggi, 
Hutan Simpan Panti, 01° 51’ 65.6”; 103° 54 10.7” 28, 19 Apr 2010, PC. Boyce, 
Siti Nurfazila Abdul Rahman & Ooi Im Hin AR- 3046 (KEP). For Borneo 
specimens see Boyce (2001). 


Distribution: West Malaysia (Johor), Borneo (widespread throughout the N 
and W, but much under-sampled). 


Habitat: Primary to secondary moist lowland to hill dipterocarp forest on 
clay and sandstone, in West Malaysia in kerapah and the drier (raised podzol) 
facies of seasonally inundated peat swamp forest; 20-840 m altitude. 


Notes: The juvenile and pre-adult stages of Rhaphidophora latevaginata and 
R. korthalsii Schott are superficially similar and to non-specialist difficult to 
differentiate. The most readily observable characters concern the petiolar 
sheath, which in R. /atevaginata is long-persistent (vs very swiftly marcescent), 
mostly adnate (vs mostly free) with the wings strongly unequal (vs weakly or 
not at all unequal), and shortly free-auriculate (vs long, narrowly triangular 
free-ligulate). Additionally, the petiole of R. latevaginata proportionately 
longer (petiole:lamina ca. 1:5 vs ca. 1:12). See Plate 1A & B. 

Pre-adult climbing stages of R. latevaginata and R. korthalsii are 
also similar but aside from the petiolar sheath characters noted above are 
readily distinguished by the feeding roots which are minutely pubescent in 
R. latevaginata and conspicuously ramentaceous in R. korthalsii. See Plate 
IC & D. 

While abundant juvenile plants were located, often intermixed with 
those of Rhaphidophora korthalsii, but only a single pre-adult, and no adult, 
plants were located during the 4-days fieldwork; by contrast, pre-adult 
and adult plants of R. korthalsii were frequent. Nonetheless we have no 
doubt that the Bornean and West Malaysian plants are one and the same 
species. Adult and fertile details given above are furnished from Bornean 
collections. 


Studies on Monstereae (Araceae) of Peninsular Malaysia II 7 


Phytogeographical Implications 


The discovery of another hitherto Bornean endemic in Peninsular Malaysia, 
following from the recent finding of Rhaphidophora megasperma in Perak 
(Baharuddin & Boyce, in press), gives further weight to the existence of a 
‘Riau Pocket’ phytochore (Corner, 1960; Ashton, 2005). 


Of particular interest is the growing evidence that the Riau Pocket 
phytochore involves plants other than trees, on which distribution and 
relationships the Riau Pocket was originally postulated. Of yet further 
interest is that while R. /atevaginata occurs in the E coast of Riau Pocket 
phytochore fragment, R. megasperma is so far known only from the Perak 
phytochore fragment. 


Figure 1. The Riau Pocket phytochore of West Malesia (areas enclosed within heavy lines 
on northwestern Borneo, Peninsular Malaysia, and Central Sumatra). From: P.S. Ashton. 
2005. Lambir’s Forest: The World’s Most Diverse Known Tree Assemblage? p. 199, Fig. 17.6. 
Used with permission. 


Acknowledgements 


This work is funded under the Research University Grant of Universiti 
Sains Malaysia (1001/PBIOLOGI/811132). 


8 Gard. Bull. Singapore 62 (1) 2010 


References 


Ashton, P.S. 2005. Lambir’s Forest: The World’s Most Diverse Known Tree 
Assemblage? pp. 191-216. In: D.W. Roubik, S.Sakai & A.A. Hamid 
Karim (eds.). Pollination Ecology and the Rain Forest: Sarawak Studies. 
Springer, New York. 


Baharuddin, S. and P.C. Boyce. 2010. Studies on Monstereae (Araceae) of 
Peninsular Malaysia I: Rhaphidophora megasperma, a new record for 
West Malaysia. Tropical Life Sciences Research: in press. 


Boyce,P.C.1999.The genus Rhaphidophora Hassk.(Araceae-Monsteroideae- 
Monstereae) in Peninsular Malaysia, and Singapore. Gardens Bulletin 
Singapore 51: 183-256. 


Boyce,P.C.2001.The genus Rhaphidophora Hassk.(Araceae-Monsteroideae- 
-Monstereae) in Borneo. Gardens Bulletin Singapore 53: 19-74. 


Corner, E.J.H. 1960. The Malayan Flora, pp. 21-24. In: R.D. Purchon 
(ed.). Proceedings of the Centenary and Bicentenary Congress of Biology. 
Singapore. 


Gardens’ Bulletin Singapore 62 (1): 9-15. 2010 9 


Studies on Monstereae (Araceae) of 
Peninsular Malaysia ITI: 
Scindapsus lucens, a New Record for Malaysia, and a Key 
to Peninsular Malaysian Scindapsus 


AHMAD SOFIMAN BIN OTHMAN, PETER C. BOYCE* 
AND CHAN LAI KENG 


Pusat Pengajian Sains Kajihayat 
[School of Biological Sciences] 
Universiti Sains Malaysia 
11800 USM, Pulau Pinang, Malaysia 
*Corresponding author: phymatarum@gmail.com 


Abstract 


Scindapsus lucens Bogner & P.C.Boyce is a species of considerable horti- 
cultural potential, arguably rivalling the commercially important S. pictus, 
originally describedfromcultivated materialofunknownprovenanceandonly 
later found wild in Sumatera. It has recently been discovered and collected 
from several localities in southern Peninsular Malaysia, representing both a 
new record for Malaysia, and through clonal propagation via various tissue 
culture techniques would supply a sustainable potential source of a local 
commercial ornamental horticultural product. An enlarged description of 
S. lucens, and a key to the Peninsular Malaysian Scindapsus are offered. A 
plate illustrating the diagnostic characters of S. /ucens is given. 


Introduction 


Scindapsus has not been revised in its entirety since 1908 (Engler & Krause, 
1908), and not treated for Malaysia since Ridley’s accounts for the flora 
(Ridley, 1907, 1925), when he recognized 5 species: Scindapsus beccarii Engl., 
S. hederaceus Migq., S. perakensis Hook.f., S. pictus Hassk., and S. scortechinii 
Hook.f. Since Ridley, fieldwork has revealed two further species: S. treubii 
Engl.,a widespread species from Sumatera to Borneo, and S. /ucens reported 
here. 

Scindapsus lucens Bogner & P.C.Boyce (1994) was described based on 
material of unknown origin cultivated in the Botanischer Garten Miinchen, 
Germany, with a note added late in proof that it had subsequently been 
located in KabupatenTapanuli, western Sumatera Utara. Since publication, 


10 Gard. Bull. Singapore 62 (1) 2010 


consultation of all the major herbaria in Peninsular Malaysia, and other 
herbaria with significant Malaysian and Indonesian collections (K, L, SING) 
has not revealed a single collection of this distinctive and horticulturally 
promising species. It was thus with some considerable surprise that during 
a recent series of fieldwork in Peninsular Malaysia S. /ucens proved to be a 
widespread species in Johor Bahru, Melaka, Negeri Sembilan, and Pahang. 
Plants seemingly favour the drier facies of perhumid podzolic shady lowland 
peatforest, although on occasions occurring with no less vigour in lowland 
mixed dipterocarp forest. 

These new collections have revealed additional morphological details 
that were unavailable at the time of original publication, and have also 
shown that the leaf colour is more variable than was initially reported, with 
the leaf blade ranging from pale grey (as per the typical collection) to an 
unusual (for Araceae) shade of jade green. 


Key to Peninsular Malaysian Scindapsus 


1. Short-climbing to perching epiphyte or scandentsemi-terrestrial............. 2 
PANAMA CS oa ces scene iaeecasetss dees Sete bance eRe 2 es eR 3 
2. Stout, short-climbing to perching epiphyte; leaves large, up to 75 x 25 cm, 


carried in litter-trapping fan, glossy deep-green, oblanceolate. Spathe white. 
Teo Watt USix. ssesacavczpicactaceseentiteueeteceanevaneren thet tases ee S. beccarii 
2. Scandent semi-terrestrial, leaves small, up to 15 x 9 cm, scattered along 
a sprawling stem, medium green, slightly glossy, ovate. Spathe yellow. 
Highlands. :scccncontie utero eee ee S. scortechinii 


(oS) 


. Leaf blade oblong lanceolate, primary lateral veins impressed adaxially 
and raised abaxially; petiole broadly conspicuously open-winged to the 
| 101) 6p bb Lan Rr eR ear RE gion oe erccecer Acree cisaasustocedocecbocolecdoaconacc-o + 

3. Leaf blade mostly cordiforme, if oblong-lanceolate then primary lateral 

veins irtually invisible; petiole not or only very obscurely winged to the 


yO) 01 kA 01 Uk nee SSE ne nor ne ere en CoC H oon sce accor oceecc nr Sacoestonsrenase 5 
A Spadix at anthesis ca S:stemyl om Pees ee eer S. hederaceus 
A Spadix at anthesis 1O=1S cml omg seeee cece eee eee S. perakensis 


5. Stems smooth; leaf blade sub-succulent, markedly oblique, oblongo-lanceolate, 
concolorous deep green, less often with jagged grey markings in juvenile 
0) £201 RRP ern RP ee crere noc ta) cccconccdcbsocodsbaueudeoseooaeoc! S. treubii 

. Stems scabrid or verruculose; leaf blade coriaceous but not markedly 
succulent, not markedly oblique, condifoniness see enna eee 6 


Nn 


Studies on Monstereae (Araceae) of Peninsular Malaysia III 1] 


6. Leaf blade conspicuously bullate, glossy pale grey to pale jade green; stems 
Vermuculose, O1GEL POLUONS TEMAIMING PTEEMN.«..<......s2:s.cce0scse0eeseec: S. lucens 
6. Leaf blade smooth, adaxially matte to somewhat scintillating, medium 
green, usually with silvery jagged spots or with these coalescing into larger 
areas; Stems at most scabrid, older portions with conspicuous orange-brown, 
EUTUE LS SPDT (SIC ce ae eee eee S. pictus 


Scindapsus lucens Bogner & P.C.Boyce, Kew Bull. 49: 789 (1994). — Type: 
Origin unknown, cultivated in Botanischer Garten Miinchen, Bogner 2113 
(holo,M!; iso,B!, K! (spirit)). Plate 1. 


Slender, sparsely branched trunk climber to ca 4 m; primary axis sterile, 
physiognomically monopodial, comprising superposed sympodia; lateral 
branches fertile, rather few and short in wild plants, producing sub-terminai 
continuation shoots after flowering. Stem up to 4mm diam., terete, epidermis 
markedly verruculate, glossy medium-green, verrucate similarly coloured, 
internodes of primary axis 3-8 cm long with one or two roots at each node; 
roots 1-2 mm diam., with grey, spongy epidermis. Leaves of primary axis 
rather distant, very regularly arranged, appressed to the substrate, those of 
lateral branches, somewhat densely grouped, petiole terete, adaxial surface 
slightly canaliculate; 3-5.5 cm x 2-3 mm, smooth, greyish to light green; 
pulvinate apically, pulvinus 1-1.5 cm, usually visible only on older leaves 
when the blade has moved its position relative to light; petiolar sheath 
2-5 cm, almost reaching pulvinus; leaf blade 7-14 x 5-9.5 cm, cordiforme, 
conspicuously bullate, coriaceous, base cordate, posterior lobes sometimes 
overlapping, apex cuspidate to acuminate, margin entire, narrowly hyaline, 
lamina adaxially glossy greyish to jade green with the raised parts of bullae 
often paler, or in green forms tinged grey, paler and glossy abaxially; primary 
lateral veins 5-7 on each side of the middle vein, ascending towards apex: 
interprimary veins barely less prominent; secondary and tertiary venation 
reticulate, much less conspicuous. Inflorescence solitary on the tips of lateral 
branches: peduncle 5 cm x 4 mm diam., terete, green; subtending cataphyll 
with very small lamina, ca 1-2 cm x 5 mm, otherwise resembling petiole; 
spathe caducous during male anthesis, 5 x 15 cm, coriaceous, cuspidate, 
exceeding spadix, somewhat constricted above spadix apex and thence 
rostrate, very pale yellow, apical rostrum ca 7 mm long, pale green prior 
to anthesis, all except the very tip becoming same colour as spathe during 
anthesis; spadix ca 2.7 x 1 cm diam., oblong-ellipsoid, fusiform, pale yellow, 
shortly stipitate, stipe ca 2 mm long. Flowers bisexual, naked, truncate; 
gynoecium rhombohexagonal in plan view, 3.5-4.2 x ca 2 mm; stylar region 
broader than ovary, ca 1 mm thick, pale yellow; stigma longitudinal-linear, 
1- 1.8 mm long, brown; ovary ca 3 mm diam., unilocular, locule globular, 


1 Gard. Bull. Singapore 62 (1) 2010 


ca 1.2 mm diam.; ovule solitary, ca 1 mm long, funicle short, placentation 
basal; stamens shorter than gynoecium, not extending markedly at pistillate 
anthesis; filaments flat, ca 1 x 1-1.2 mm; thecae ellipsoid, ca 0.8 x 0.5 mm, 
opening by a subapical slit. Infructescence unknown. 


Specimens seen: MALAYSIA, Johor Bahru: Mersing, Kluang— Mersing road, 
km 39, Lenggor FR.,02° 15.727’, 103 43.767’, 18 April 2010, PC. Boyce, Siti 
Nurfazilah bt Abdul Rahman & Ooi Im Hin AR-3056 (KEP); Kota Tinggi, 
Panti FR.,01° 52.226’, 103 54.755’, 19 April 2010, PC. Boyce, Siti Nurfazilah 
bt Abdul Rahman & Ooi Im Hin AR-3057 (KEP); Kota Tinggi, Hutan Lipur 
Panti; 01° 48.077’, 103° 57.202’, 19 April 201, PC. Boyce, Siti Nurfazilah bt 
Abdul Rahman & Ooi Im Hin AR-3060 (KEP). Melaka: Machap, Hutan 
Simpan Bukit Sedana, 02° 24’ 00.6”; 102° 20° 91.7”, 11 May 2010, PC Boyce 
& Ng Kiaw Kiaw AR-3059 (KEP); Negeri Sembilan: Pasoh Forest Reserve, 
3° 0’ 0; 102° 19° 60, 13 May 2010, PC. Boyce & Ng Kiaw Kiaw AR-3058(KEP); 
Pahang: Kuala Rompin, Taman Negeri Endau Rompin, Waterfall trail, 02° 
37° 08.9”; 103°20° 81.3”, 12 May 2010, PC. Boyce & Ng Kiaw Kiaw AR-3063 
(KEP); Jerantut, Krau Wildlife Centre, main trail to river, 03° 49° 39.2”; 
102°13’ 03.1”, 13 May 2010 PC. Boyce & Ng Kiaw Kiaw AR-3054 (KEP). 


Distribution: Malaysia (Johor Bahru, Melaka, Negeri Sembilan & Pahang). 
Indonesia, (western Sumatera Utara). 


Habitat: Drier facies of perhumid lowland shady peatforest; less often in 
lowland mixed dipterocarp forest (e.g., at Pasoh FR., Krau W.R.). 20-125 
masl. 


Notes: Scindpasus lucens appears to be a widespread species in southern 
Peninsular Malaysia, with occurrences so far recorded in four states. 
Curiously, given the complete absence of herbarium specimens, it appears 
to be more abundant than S. pictus, despite the abundance of material of the 
latter species in herbaria. 


Potential of Scindapsus lucens as a sustainably-produced 
ornamental horticulture subject 


Scindapsus lucens is a widespread species in the southern Peninsular, and 
although never abundant appears to be more prevalent than S. pictus 
Hassk., the species to which S. /ucens is most similar. The very attractive 


Studies on Monstereae (Araceae) of Peninsular Malaysia III 


Plate 1. Scindapsus lucens Bogner & P.C.Boyce. A. Juvenile primary axis. Note the glossy 
quality of the leaf blade (PC. Boyce & Ng Kiaw Kiaw A R-3054). B. Juvenile plant, jade-green 
type.(P.C. Boyce & Ng Kiaw Kiaw AR-3063). C. Type clone in cultivation, Jardin Botanique 
du Montet, Nancy, France (Bogner 23/1). D. Detail of stem showing the diagnostic verrucate 
surface (P.C.Boyce & Ng Kiaw Kiaw AR-3054). Images A & B, D © P.C.Boyce; Image C © 
David Scherberich, used with permission. 


14 Gard. Bull. Singapore 62 (1) 2010 


Plate 2. A. Scindapsus treubii Engl. Note the subsucculent, strongly oblique leaf blade, and 
smooth stem. B-E. Scindapsus pictus Hassk. B. Primary axis showing the diagnostic and 
older portions with conspicuous orange, brittle scabrid epidermis. C-E. A selection of leaf 
markings typical of S. pictus. Notes the somewhat scintillating quality particulary in C & E. 
Images © P.C. Boyce. 


Studies on Monstereae (Araceae) of Peninsular Malaysia II] 15 


leaves and, for a lianescent aroid, small habit make S. /ucens an ideal subject 
for consideration as a commercial ornamental product with via various in 
vitro culture technology. The ease with which S. pictus is globally produced 
in tens of millions for the ornamental horticulture market augers well that 
S. lucens should also prove similarly amenable. In many respects S. /ucens 
is a far more attractive plant than S. pictus, not in the least because it has a 
much more compact habit, and is not a heteroblastic species and thus the 
attractive juvenile stage is retained into adult-hood, unlike that situation 
with S. pictus, which requires regular pruning to retain the commercially 
desirable leaf form of the juvenile. Since S. /ucens has a high potential as 
an attractive ornamental plant, it may be over-collected by irresponsible 
individuals to fulfil the public demand for new ornamental plant in the 
market. This may lead to the eventual extinction of this plant. To conserve 
the germplasm and prevent over-collection from its natural habitat, S. /ucens 
can be mass propagated via induction of multiple shoots formation using 
shoot culture technique or the production of somatic embryogenesis using 
the leaf as explants. Millions of plantlets can then be produced to fulfil the 
market demand. 


Acknowledgements 


This work is funded under the Research University Grant of Universiti 
Sains Malaysia (1001/PBIOLOGI/811132) 


References 
Boyce,P.C.1999.The genus Rhaphidophora Hassk.( Araceae-Monsteroideae- 
Monstereae) in Peninsular Malaysia, and Singapore. Gardens’ Bulletin 


Singapore 51: 183-256. 


Engler, A. & K. Krause. 1908. Araceae — Monsteroideae. In: A. Engler (ed.), 
Das Pflanzenreich 37(1V.23B): 4-139. 


Ridley, H.N. 1907. Araceae. Materials for a Flora of Malay Peninsula. 3: 1-53. 


Ridley, H.N. 1925. The Flora of the Malay Peninsula, 5: 84-131. Reeve & Co., 
London. 


Gardens’ Bulletin Singapore 62 (1): 17-22. 2010 17 


Begonia droopiae Ardi (Begoniaceae), 
a New Species of Begonia 
from West Sumatra 


W.H.ARDI AND M. HUGHES 


Bogor Botanic Garden 
Jl. Ir. H. Juanda No. 13, P.O. Box 309, Bogor, Indonesia 
- Royal Botanic Garden Edinburgh 
20A Inverleith Row, Edinburgh EH3 SLR, U.K. 
E-mail: m.hughes@rbge.ac.uk 
Author for correspondence: wisn001 @lipi.go.id 


Abstract 


A new species of Begonia (Begoniaceae), B. droopiae Ardi,is described from 
the Indonesian island of Sumatra. It belongs to Begonia sect. Reichenheimia 
and is a limestone endemic in the Sawah Lunto District. Its IUCN threatened 
category is considered to be ‘Vulnerable’. 


Introduction 


During an expedition to West Sumatra in August 2009 organised by The 
Royal Botanic Garden Edinburgh, Kebun Raya Bogor and Andalas 
University Herbarium (ANDA), a new species of Begonia, B. droopiae 
Ardi, was collected from a limestone cave in the Batang Pangean I Nature 
Reserve, Sawah Lunto District, West Sumatra Province. Begonia droopiae 
is placed in Begonia sect. Reichenheimia because it exhibits the typical 
characters of the section: rhizomatous habit, protandrous inflorescences and 
three locular fruit with entire placentae. 

This species-rich genus was previously represented by 52 species in 
Sumatra (Hughes, 2008; Hughes er a/. 2009), although it is obvious from 
herbarium collections that many more remain to be described from the 
island. Begonia sect. Reichenheimia is the second-most species rich section 
of the genus in Sumatra, with 11 species. All available herbarium specimens 
in ANDA, BO, E and SING have been consulted, and hence it must be 
assumed, at least until more intensive collecting in West Sumatra may reveal 
otherwise, that this species has a very restricted range (Fig.1). 


18 Gard. Bull. Singapore 62 (1) 2010 


100°0'0"E 1050'0"E 


00'0" 


50'0" 


95°0'0"E 100°0'0"E 


Figure 1. Distribution of Begonia droopiae. 


Begonia droopiae Ardi, sp. nov. (Sect. Reichenhemia) 

Haec species Begoniae nurii similis est sed foliis magis asymmetricis, floribus 
femineis tepalis 3 (nec 2) provisis et stipulis secus costam pilosis recedit. 
—Typus: Indonesia, Sumatra, West Sumatra (Sumatra Barat), Batang Pangean 
Nature Reserve, Sawah Lunto District, Nagari Solok Ambah, Perkaulan 
cave, 00° 43’ 21.7” S, 101° 09’ 01.0” E, 484 m, 21 viii2009, A. J. Droop, W. H. 
Ardi, Nurainas & Riki AJD173 (holo, BO; iso, E, BO, ANDA). Plate 1. 


Perennial, creeping, monoecious herb, to ca 10 cm tall, rooting at the nodes, 
hairy with up to ca 2 mm long, multicellular hairs. Stems rhizomatous, 
internodes very short, up to ca 2 mm long, with scattered multicellular 
hairs; stipules 3-6 x 3-5 mm, triangular, with an abaxially prominent midrib 
forming an up to ca 4 mm long, thin, hairy appendage at the apex, abaxially 
densely hairy along the midvein, persistent. Leaves alternate; petioles 5- 
19.5 cm long, sparsely to densely hairy; lamina basifixed, 3.5-9.5 x 2-6.8 cm, 
very asymmetric, ovate to elliptic, base cordate, lobes sometime slightly 
overlapping, apex acuminate, margin broadly crenate and fringed with 
hairs, adaxial surface glabrous, purplish-green to dark purple between the 


Begonia droopiae Ardi (Begoniaceae) from West Sumatra 19 


“>N ra 
ee 


«% 


Plate 1. Begonia droopiae Ardi. A. Habit; B. Stipules (scale bar = 5 mm); C. Male flowers 
(scale bar = 5 mm); D. Female flower (scale bar = 5 mm); E. Fruits (scale bar = 5 mm). A-E 
based on A. J. Droop, W. H. Ardi, Nurainas & Riki AJD173. 


20 Gard. Bull. Singapore 62 (1) 2010 


veins, slightly raised, veins green, abaxial surface pale green to dark purple 
between the veins, hairy on the veins, venation palmate. Inflorescences 
cymose, axillary, protandrous, bisexual; peduncles 5-11 cm long, reddish, 
glabrescent to sparsely hairy; bracts sub orbicular, ca 1.5-2.5 x 1-2 mm, 
margin fimbriate, deciduous. Male flowers, pedicels 8-20 mm, moderately 
hairy; tepals 4, two outer tepals, white or white with a tinge of pink, 11-17 
x 6-7 mm, elliptic to sub orbicular, base slightly cordate, apex rounded, 
abaxially moderately red hairy; two inner tepals white, 7-9.5 x 3-3.5 mm, 
oblong-obovate, glabrous; androecium of ca 40 stamens, yellow, filaments ca 
1 mm long, fused at the base, anthers ca 1-1.5 mm long, obovate, dehiscing 
through unilaterally positioned slits ca 1/2 as long as the anther, connective 
not projecting. Female flowers, pedicels 6-9 mm, glabrous; tepals 3, unequal, 
two outer tepals orbicular to sub orbicular, 5.5-6 x 4.5-6 mm, one inner 
tepal, obovate, 5-6 x 1.4-1.6 mm; ovary 6-7 x 10-13 mm, orbicular, locules 3, 
placentation axile, placentae entire; wings 3, subequal, triangular, rounded at 
the and the apex, the widest point at the middle of the ovary, glabrous; style 
basally fused, 3-branched, each stylodium bifurcate in the stigmatic region, 
stigmatic surface a spirally twisted papillose band, yellow. Fruits, on thin, 6-9 
mm long, sparsely hairy pedicels; capsules 8-8.5 x 10-7 mm, (excluding the 
wing) deflexed, dehiscent, splitting along the wing attachment, drying pale 
brown, glabrous, wings 4-5 mm wide at the widest point (at the middle of the 
ovary), wing shape as for ovary; seeds unknown. 


Distribution: Indonesia, Sumatra, West Sumatra (Sumatra Barat), Batang 
pangean nature reserve, Sawah Lunto District, Nagari Solok Ambah, 
Perkaulan cave. 


Habitat: This species is growing directly on vertical limestone wall in the 
front of cave at 484 m altitude. 


IUCN Conservation category: We consider this species to belong to the 
VUD2 IUCN category, as it has a very restricted distribution. Although it 
resides in a protected area, there is some evidence of small scale agricultural 
activities encroaching the habitat nearby. 


Notes: Begonia droopiae is morphologically similar to Begonia nurii Irmsch. 
Both species exhibit a rhizomatous habit, patterned leaves, and have male 
flowers with 4 tepals. However, B. droopiae can easily be distinguished 
from B. nurii by its very asymmetric leaves with an acuminate apex, and 
stipules which are hairy on the mid-vein (versus slightly asymmetric leaves, 
with a rounded apex and stipules which are glabrous on the mid-vein in B. 
nurit). The tepal number of the female flowers also differs; B. droopiae has 


Begonia droopiae Ardi (Begoniaceae) from West Sumatra 21 


three tepals, whilst B. nurii has two. The wing shape of the fruits is a further 
difference: in B. droopiae the wings are rounded at the base and the apex, 
whilst in B. nurii the wings are rounded at the base and cuneate at the apex. 
Begonia rajah Ridlis another allied species from Peninsular Malaysia, similar 
in its rhizomatous habit, patterned leaves and male flowers with 4 tepals (R. 
Kiew, pers. comm.), but B. droopiae differs in having very oblique leaves 
(the midrib is clearly at an acute angle) which are soft and not succulent; B. 
rajah has thickly succulent leaves which are more bullate,1.e., raised between 
the veins. Further, Begonia rajah has never been found on limestone; most 
Peninsular Malaysia Begonia species grow either on limestone and not on 
other rock types or vice versa (Kiew, 2005). A comparison of the two species 
to Begonia droopiae is presented in Table 1. The epithet is after the first 
collector, Alison Jane Droop. 


Table 1. Comparison of Begonia droopiae, B. rajah and B. nurii. 


Begonia Begonia Begonia 
droopiae rajah nurii 


Asymmetric Slightly asymmetric 
Thickly succulent Thin 


3.5-9.5 x 2-6.8 cm 7-15 x 6-15 cm 2-7 x 3-11 cm 


: Cordate and Cordate and slightly Cordate and often 
Lamina base : ; : , 
slightly overlapping overlapping overlapping 
Lamina apex 


Surface between 
the veins 


Character 


Acuminate Short and acute Rounded 


Slightly prominent Prominently raised Slightly prominent 


eee 


Hairy on mid vein Midvein glabrous 


Stipules Midvein glabrous 


Peduncle 5-11 cm long 10-25 cm long 4-19 cm long 


8-8.5 x 10-12 mm 6-7 x 5-6 mm 5-9 x 10-16 mm 


Fruit size 


E th 3 
: Rounded at the Romadediat the base Rounded at the base, 
Wing shape sub truncate at the 
base and the apex cuneate at the apex 


apex 


i) 
i) 


Gard. Bull. Singapore 62 (1) 2010 


Acknowledgements 


We are grateful to the Indonesian Ministry of Research and Technology 
(RISTEK), the Indonesian Institute of Sciences (LIPI), Direktorat Jenderal 
Perlindungan Hutan dan Konservasi Alam (DITJEN PHKA) for giving 
us permission to conduct research in West Sumatra and for their help and 
assistance. The curators of herbaria ANDA, BO, E and SING are thanked 
for facilitating access to specimens. The fieldwork was supported by the 
ASEAN Centre for Biodiversity. Prof. Ruth Kiew is gratefully acknowledged 
for her constructive comments, and Dr. Robert Mill is thanked for his Latin 
expertise. 


References 


Doorenbos, J., M.S.M. Sosef and J.J.RE. de Wilde. 1998. The sections 
of Begonia including descriptions, keys and species lists (Studies in 
Begoniaceae VI). Agricultural University Wageningen Papers 98(2): 1- 
266. 


Hughes, M. and M. Pullan. 2007. Southeast Asian Begonia Database. 
Electronic publication accessible via: www.rbge.org.uk. 


Hughes, M. 2008. An annotated checklist of Southeast Asian Begonia. Royal 
Botanic Garden Edinburgh, UK. 


Hughes, M., D. Girmansyah, W.H. Ardi and Nurainas. 2009. Seven New 
Species of Begonia from Sumatra. Gardens’ Bulletin Singapore 61(1): 29-44 


Kiew, R.2005. Begonias of Peninsular Malaysia. Natural History Publications, 
Borneo. 


Gardens Bulletin Singapore 62 (1): 23-30. 2010. 23 


Newly Recorded Endiandra R. Br. (Lauraceae) from 
Waigeo Island, Raja Ampat, Papua, Indonesia 


D. ARIFIANI 


Herbarium Bogoriense, Research Center for Biology, Indonesian Institute 
of Sciences (Pusat Penelitian Biologi-LIPI), Jl. Raya Jakarta-Bogor 
KM 46, Cibinong 16911, 
West Java, Indonesia 
E-mail: debyarifiani@yahoo.com 


Abstract 


Endiandra trees are not frequently encountered in the forest in Indonesia. 
Recent exploration in Waigeo Island resulted in the collection of three 
species of Endiandra. All three species were collected for the first time from 
Waigeo Island. The three species are restricted in distribution to eastern 
part of Indonesia (Papua Province) and Papua New Guinea. Key to the 
species and species description are provided. 


Introduction 


Waigeo Island is part of the Raja Ampat Islands, an archipelago that 
is situated at the northern extremity of Indonesia’s easternmost Papua 
province. Waigeo Island is the largest island in the archipelago, 130 km long 
and 48 km wide and positioned at the northernmost of Raja Ampat Islands. 
Raja Ampat is very rich in both terrestrial and oceanic biodiversity including 
plant, coral reefs and fish (Anonymous 2006). 

Endiandra is one of the genera in the family Lauraceae, a dominant 
plant family in tropical forests and occurs from lowland up to montane 
forests. Main distribution area of Lauraceae species is in the tropical forests 
of South East Asia and America (Heywood, 1993). 

Endiandra was first described by R. Brown (1810) based on the 
type species, Endiandra glauca, from Australia. The genus consists of 
approximately 100 species distributed from South China, Taiwan, Malesia, 
and Australia up to Fiji (Rohwer, 1993). In Malesia, New Guinea is the 
main distribution centre of Endiandra with several endemic species. A 
floristic survey to Waigeo Island was carried out to study the occurrence of 
Endiandra on the island and to make additional collection for the Herbarium 
Bogoriense. 


24 Gard. Bull. Singapore 62 (1) 2010 


Materials and Method 


Herbarium specimens studied for this research were collected 
from Waigeo Island. Additional specimens of Endiandra available at the 
Herbarium Bogoriense were also examined. Main localities explored on 
Waigeo Island are forests in the area of Teluk Mayalibit District and riverine 
forest along the Werabiyai River. 

We collected vegetative, floral and fruiting parts of each plant species 
and took their pictures. Information, such as locality, plant habit, uses, color 
and scent of specimens, were noted to help make the species identification 
easier. Identification was done mainly by consulting references and 
comparing the recently collected herbarium with the herbarium available 
at the Herbarium Bogoriense. The descriptions for each species were 
constructed based on observed morphological characters of the collected 
specimens. 


Results and Notes 


The forests at the Waigeo Island are mostly of moist lowland type. The 
areas of Warsamdim and Werabiyai on the island vary from slightly hilly to 
steep slope. Easily accessible forests were disturbed, especially by logging 
activities in the past. However, good vegetation can still be found on hills 
and slopes. 

Endiandra species are less known because they are not easily found 
in the forests in general, neither in Waigeo forests. The more common 
genera of Lauraceae in the area are Actinodaphne, Cryptocarya and Litsea, 
of which young trees are easily spotted inside the forests. However, it is 
very fortunate that in the recent floristic exploration in Waigeo Island, we 
encountered three species of Endiandra. 

Three species of Endiandra, i.e., Endiandra beccariana, E. grandifolia 
and EF. papuana, were collected from the island. 


Endiandra R. Br. (Lauraceae) 


Taxonomically, the genus Endiandra is characterized by having alternate, 
spiral and pinnately-veined leaves, inflorescences paniculate-determinate 
with ultimate flowers not strictly opposite. Flowers trimerous, bisexual, 
stamens 3, each with 2 thecae, glands present or absent. In some species 
glands are united forming a disc surrounding the stamens and pistil. Fruits 
are without cupule, free on receptacle. 


Newly Recorded Endiandra R. Br. (Lauraceae) 25 


The genus Endiandra is known for its wood. E. palmerstonii produces 
high quality wood commonly used for good furniture and for building 
construction. 


Identification key to species of Endiandra in Waigeo Island 


1. Lamina broadly elliptic, lateral veins 11-15 pairs, glands reniform, not united 


SRPMS PANICAIS PES. os ee 3 eee ctcs-s.sesersersesesssseees 2. EB. grandifolia 
1.Lamina elliptic, lateral veins 7-10 pairs, glands united forming disc-like 
2 UL EL LEESS cc this Sok eee eee 2. 
2. Leaf surfaces and panicles sparsely pubescent ............. 1. E. beccariana 
2. Leaf surfaces and panicles densely pubescent ................ 3. E. papuana 


Species description 


1. Endiandra beccariana Kosterm., Reinwardtia 7: 5 (1969) 474. —Type: 
Morotai Isl., District Tobelo, N. Totodoku, 14 May 1949, Kostermans & 
Tangkilisan 146 (A, BO). Plate 1. 


Tree up to 32 m, 40 cm dbh. Bark greyish brown, lenticellate. Young twigs 
dense erect pubescent, rusty colored. Terminal buds densely pubescent, 
rusty colored, 8 mm long and 3 mm wide. Leaves spiral, stiffly chartaceous, 
elliptic; 8.5-19 x 6-11 cm; apex acuminate, base cuneate or slightly rounded: 
both surfaces finely reticulate; upper surfaces shiny, midrib flat, lateral 
veins slightly impressed; lower surfaces slightly shiny, sparsely pilose or 
glabrescent, midrib raised; lateral veins 7-10 pairs, prominent below, arcuate 
towards margin; petiole terete, densely pubescent to glabrescent, flat, 12- 
23 x 10-20 mm. Inflorescences paniculate, axillary, 4-11 cm long, pubescent. 
Pedicel slender, ca 3 mm long. Flowers yellow, ca 6 mm in diameter; tepals 
fleshy, spreading, narrowly elliptic, slightly acuminate, 2.5-3 mm long; glands 
united in a form of a disc, ca 0.75 mm thick, brownish yellow; anthers widely 
triangular, emerge from gland-like disc; locules large; ovary ellipsoid, 
glabrous; style 0.5 mm long, stigma unconspicous. Fruit ellipsoid, ca 1-2 cm 
long, green, free on the receptacle. 


Recently collected specimens: INDONESIA. Papua Province: Kabupaten 
Raja Ampat, Waigeo Isl., District Teluk Mayalibit, Desa Warsamdim, 120 
m, 8 Jun 2007, D. Arifiani & Obaja 596 (BO); ibid., + 50 m, 13 Jun 2007, D. 
Arifiani & Obaja 652 (BO). 


26 Gard. Bull. Singapore 62 (1) 2010 


Additional specimens examined: INDONESIA. Sorong, Kadamah, 14 Aug 
1948, Main 592 (BO); Manokwari, Warnapi 15 km N. of Ransiki, 10 m, 25 
Sep 1948, Kostermans 448 (BO); Manokwari, Warnapi, 20 m, 30 Sep 1948, 
Kostermans 491 (BO); Morotai Isl., along Sambiki R. (S.E. Morotai), 30 m, 
22 May 1949, Kostermans 854 (BO); Morotai Isl., along Sambiki R., 100 m, 
22 May 1949, Kostermans 890 (BO); Morotai Isl.,23 May 1951, Kostermans 
934 (BO); N. slopes of upper Aifat Valley, between Senopi and Aifatfekaan, 
W. of Kebar Valley, 10 Dec 1961, Moll BW 12908 (BO). 


Note: Endiandra beccariana was first described by Kostermans in 1969 based 
on specimens from Morotai Island, Manokwari, Sorong, and Ramoi. In our 
exploration, we spotted the species in two locations on Waigeo Island. 

This species is similar to the E. papuana Lauterbach, which both can 
only be differentiated by checking the amount of indument on the lower leaf 
surface and the inflorescences. Kostermans (1969) noted that the panicles 
of E. beccariana are glabrous and I would not agree with his statement given 
the fact that the panicles bear sparsely erect indument. 

The specimens of EF. beccariana collected from this floristic survey were 
anew record of the species occurrence on Waigeo Island. Known previously 
from Morotai Island and Northwest part of West Papua, its occurrence 
in Waigeo Island indicates a possible connection by seed dispersal from 
Morotai Island to Papua and/or vice versa (Fig. 2). 


2. Endiandra grandifolia Teschn., Engl. Bot. Jahrb. 58 (1923) 417. —Type: 
Papua New Guinea, 10 May 1909, Schlechter 17691 (K). 


Tree up to 25 m high, 45 cm dbh. Twigs hairy. Leaves simple, coriaceous; 
broadly elliptic; 14-32 x 10-17 cm apex acute to rounded; base obtuse; 
both surfaces finely reticulate, upper surfaces shiny, midrib slightly raised, 
lateral veins slightly impressed, with indument; lower surfaces densely 
tomentose; midrib raised below; lateral veins 11-15 pairs, prominent below, 
arcuate towards margin; petiole 1.5 cm long, stiff, tomentose. Inflorescences 
paniculate, compact, up to 9 cm long, axillary, densely tomentose, reddish 
brown, many flowers. Tepals slightly unequal; elliptic; apex obtuse; tomentose. 
Stamens elliptic, tip rounded, pubescent; glands reniform, stalked, pilose; 
staminodia cordate, pilose, stalked. Ovary ovoid, glabrous. Fruit unknown. 


Recently collected specimens: INDONESIA. Papua Province: Kabupaten 
Raja Ampat, Waigeo Isl., District Teluk Mayalibit, Desa Warsamdim, low 
alt., Jun 2007, Mirmanto & Ruskandi 09305 (BO). 


ona 


“s 


; 
' 
. 


Plate 1. Endiandra beccariana: A. Tree habit (D. Arifiani & Obaja 652); B. Leaf arrangement: 
C. Flower buds; D. Mature flower, glands united forming disc-like appendages with stamens 
exposed from the middle: E. Young fruit. (Photos C-E: A. Hidayat). 


28 Gard. Bull. Singapore 62 (1) 2010 


Additional specimens examined: PAPUA NEW GUINEA. Madang District, 
Terr. of New Guinea, near the Gogol River by Mawan village (ca 25 km 
inland), 60 m, 22 Jun 1955, Hoogland 4919 (BO); Morobe District, Lae 
Subdistrict, Oomsis Logging area N.G.I., 180 m, Millar NGF 12036 (BO); 
Morobe District, Menyamya Subdistrict, between Aseki & Menyamya, 
Spreader Divide, 21 Nov 1970, Streimann & Kairo NGF 42451 (BO). 


Note: E. grandifolia may be differentiated from both E. beccariana and 
E. papuana by observing the shape or fusion of the glands. Some species 
of Endiandra, including E. grandifolia, bear three pairs of glands at the 
bases of each stamen. However, these glands sometimes may unite to each 
other because of the limited space inside the flower and forming a disc-like 
appendage surrounding the stamens. This is the case with both E. beccariana 
and E. papuana. 

This survey yielded the first collection of Endiandra grandifolia 
from western part of New Guinea. Previously, the species was only collected 
from Madang and Morobe districts of Papua New Guinea. Occurrence of 
E. grandifolia in Waigeo Island extend its distribution toward northwestern 
part of New Guinea (Fig. 2). 


3. Endiandra papuana Lauterbach, Nova Guinea, Bot. 8: 2 (1912) 819. - 
Type: Papua New Guinea, 6 Dec 1907, Branderhorst 263 (BO, K). 


Tree up to 30 m. Twigs densely pilose, rusty colored. Terminal buds densely 
pubescent, smooth, rusty colored. Leaves spiral, elliptic, stiffly chartaceous, 
10-19 x 7-10 cm, apex acute, base obtuse to acute; both surfaces densely 
reticulate; upper surface shiny, midrib flat, lateral veins slightly impressed; 
lower surface slightly shiny, densely pilose, midrib raised; lateral veins 7- 
8 pairs, raised, arcuate towards margin; petiole terete, densely pubescent, 
1.5-2.0 x 0.20-0.25 cm. Inflorescences paniculate, densely pubescent, up to 
18 cm long, axillary. Tepals spreading; glands large, united to form a disc- 
like appendage. Fruits spherical to ellipsoid, 1-1.5 cm long, free on the 
receptacles. 


Recently collected specimens: INDONESIA. Papua Province: Kabupaten 
Raja Ampat, Waigeo Isl., along Werabiyai River, Jun 2007, Mirmanto & 
Ruskandi 09304 (BO). 


Additional specimens examined: WEST NEW GUINEA, Albatros Bivouac, 
7 May 1926, Docters van Leeuwen 9016 (BO). PAPUA NEW GUINEA, 
Morobe District, Wareo, 450 m, 5 Feb 1936, Clemens 1782 (BO); Northern 


Newly Recorded Endiandra R. Br. (Lauraceae) 29 


Division, Terr. of Papua, between Mambare and Arumu Rivers. South of 
Botue village (near Kokoda), 350 m, 21 Sep 1953, Hoogland 3955 (BO): 
Oomsis Logging area, NW of Lae, Morobe District. T.N.G. 90m. 5 Mar 
1959, White NGF 10487, Near Garagos, Lae. Bulolo Road. Morobe District. 
450 m, 3 May 1962. Havel & Kairo NGF 11197 (BO): Titapuba, Morobe 
District: 10 Jan 1966, Streimann & Kairo NGF 26160 (BO). Tributary of 
Busu River, above Sankwep R., Lae Subdistrict, Morobe District. 13 Apr 
1972, Wommersley NGF 43919 (BO). 


Note: E. papuana 


is More commonly encountered in the forests of Papua 


New Guinea, around Morobe District than in Indonesian part of New 
Guinea. Specimen of EF. papuana collected from Waigeo Island is a good 
addition to the Herbarium Bogoriense collection (see Fig. 3). 


(far 

En? he: Raja Ampat 
ESE 

a = | . oS, 

Si =. ae A ea = 

~ ¥ cr oc - " 

Paes Ss eee Wis + i 

= ej = os : ~~ 

Py a ae a ee 
S x Sa 
re aot = — ‘ a. 

> . =< fo 4 = % 

[r~.* F fr) a 
‘ * 5 4 | 
| i et eo =o 
si ome is ~ 
i st Es = =t 
| —_ , j 
sor enn She a 
‘sal ee 4 i. TE 


Figure 2. Distribution of Endiandra beccariana (m). E. grandifolia (a) and E. papuana (e). 


: Waigeo 
= £ 
? aS =< 
ae 
Fe _ : & 
© c — 
a — 
ee oe 
ee ee 
Tu"; 
a ye 
2 5S we s 
—— rs f 
a “Ue? 3 
+ 
fae te 


Figure 3. Occurrence of newly recorded species of Endiandra in Waigeo Island: Endiandra 
beccariana (m). E. grandifolia (a) and E. papuana (@) 


30 Gard. Bull. Singapore 62 (1) 2010 


Acknowledgements 


The field research was funded by DIPA project of the Research Center for 
Biology-Indonesian Institute of Sciences (LIPI) for the fiscal year 2007/2008. 
I would like to thank Nurdin, A. Sujadi, A. Ruskandi, people of Warsamdim 
village (especially: Obaja, Andi, Yustus, Dani, Marcus, and Mattheus) for 
their valuable assistance during the field survey, Suhardjono and Roemantyo 
for assisting with distribution map. My respect goes to the late A. Ruskandi 
for his enormous knowledge of plant identification who passed away in 2008 
after his short battle with cancer. Indonesian Plant Taxonomy Association 
(Penggalang Taksonomi Tumbuhan Indonesia) and DIKTI facilitated author 
and editor meeting for manuscript preparation. 


References 


Anonymous. 2006. Atlas Sumberdaya Pesisir Kabupaten Raja Ampat, Provinsi 
_ Irian Jaya Barat. Pemerintah Kabupaten Raja Ampat dan Konsorsium 
Atlas Sumberdaya Pesisir Kabupaten Raja Ampat, Irian Jaya Barat. 


Brown, R. 1810. Laurinae. In: Prodromus Florae Novae Hollandiae et Insulae 
Van Diemen. Typis Richardi Taylor et Soci, London. 


Heywood, V. 1993. Flowering Plants of the World. B. T. Batsford Ltd. 
London. 


Kostermans, A.J.G.H. 1969. Material for a revision of Lauraceae II. 
Reinwardtia 7(5): 470-496. 


Lauterbach, C. 1912. Lauraceae. Nova Guinea 8(2): 819-820. 


Rohwer, J.G. 1993. Lauraceae. In: K. Kubitzki, J.G. Rohwer, V. Bittrich (eds.). 
The Families and Genera of Vascular Plants IT. Springer Verlag, Berlin. 


Teschner, H. 1923, Die Lauraceen Nordost-Neu-Guineas. In: A. Engler 
(ed.). Botanische Jahrbiicher fiir Systematik, Pflanzengeschichte und 
Pflanzegeographie 58: 413-420. 


Gardens Bulletin Singapore 62 (1): 31-42. 2010. 


Species of Marasmius (Agaricales: Tricholomataceae) 
from Kayan Mentarang National Park, East Kalimantan, 
Indonesia 


ATIK RETNOWATI 


Herbarium Bogoriense 
Research Center for Biology, The Indonesian Institute of Sciences 
Bogor, Indonesia 


Abstract 


Five species of Marasmius were encountered from forest surrounding 
Pa 'raye village at Kayan Mentarang National Park, East Kalimantan: three 
of them are described as new taxa (1 new species and 2 new varieties). 
The five species of Marasmius are M. cf. purpureostriatus, M. guyanensis, 
M. coklatus var. mentarangensis, var. nov., M. caryote var. pa’rayeensis, var. 
nov., and M. gypseus, sp. nov. Comprehensive descriptions, illustrations, 
and comparison with similar taxa are presented. 


Introduction 


Among the Agaricales in the tropics, Marasmius has become a favorite 
collection item. Species of Marasmius grow relatively longer period of time 
than other agarics. They are very attractive and relatively easy to be spotted. 
It is the most common genus in tropic, and grows frequently on forest soil, 
fallen leaves, and on dead or living wood and other plant tissues, such as 
grass roots and bamboo twigs. Most species of Marasmius are saprophytic, 
some are even parasitic, and none goes into mycorrhizal association. 

The earliest report of Marasmius from Indonesia with 28 species 
was made by Léveillé (1844, 1846) and many of them were reported from 
Java. This was followed by several reports by Moritzi (1845-1846), Zollinger 
(1854), Hennings (1900), Overeem and Overeem-de Haas (1922), Boedijn 
(1940). Desjardin, Retnowati, and Horak (2000) documented 37 species 
of the genus from Java and Bali. However, no species of Marasmius was 
recorded from Kalimantan, particularly from Kayan Mentarang National 
Park, East Kalimantan. This paper presents five species of Maramius 
encountered during a three weeks expedition to survey the fungal flora 
from the forests surrounding Pa’raye village at Kayan Mentarang National 
Park in April of 2003. 


ies) 
ir) 


Gard. Bull. Singapore 62 (1) 2010 


Kayan Mentarang National Park, which lies between altitudes 
116°15’-115° E and 2°-4°30’ N, embraces a total area of 1.35 million hectares. 
It is located in East Kalimantan, bordering with Sabah and Sarawak. The 
inventory was done from several collecting sites in the mountainous Pa’raye 
village located north of the National Park. Descriptions of the species are 
based on fresh collections made by the author. All micro characters were 
studied from dried materials rehydrated in distilled water and 3% of KOH 
solution, with the use of Melzer’s reagent or Congo Red dye. Color terms 
and notations are those of Kornerup and Wanscher (1978). Spore sizes were 
based on measurements of 25 basidiospores. 

Spore statistics include - X = the arithmetic mean of the spore length 
by spore width (+ standart deviation); Q, the quotient of spore length and 
spore width in any one spore to indicate the range of variation in n spores 
measured; q, the mean of Q-values in a single sample. 

The basidiomes in illustration are of natural size, basidiospore (scale 
bar = 12 um); basidia, cystidia, pileipellis and stipitipellis (scale bar = 15.7 
um). All collections examined are deposited in Herbarium Bogoriense 
(BO) and Harry D. Thiers Herbarium (SFSU). 


Description of taxa 


1. Marasmius cf. purpureostriatus Hongo 
J. Jap. Bot. 33: 344. 1958. -Type: Japan, Otsu City, Ishtyama, 7 May 1957, 
Hongo 1609 (Isotype, ZT [3221]). Fig. 1. 


Pileus 55 mm diam, convex with slightly depressed center, sulcate; margin 
crenate, straight; surface pruinose, smooth; greyish magenta (13-E3) overall. 
Context thin, pale greyish magenta. Lamellae adnate, distant (12 attached 
stipe) with 1 series of lamellulae, narrow, non-marginate; pale greyish 
magenta. Stipe 180 x 5 mm, hollow, central, equal, cylindrical; smooth, 
pruinose; pale greyish magenta. Odor and taste not distinctive. Basidiospores 
unobserved. Basidia 46.4-56 x 12-12.8 um, 4-spored, clavate. Basidioles 
clavate. Cheilocystidia common, 16-24 x 7.2-12.8 um, clavate, broadly clavate, 
sub-globose to globose, hyaline to weakly yellow, thin-walled. Pleurocystidia 
absent. Pileipellis hymeniform, clavate to broadly clavate or subglobose, 
16-25.6 x 8.8-9.6 um, non-diverticulate, hyaline, thin-walled. Stipe tissue 
monomitic; cortical and medullary hyphae indistinguishable, 6.4-20 um diam, 
parallel, cylindrical, hyaline, thin-walled, strongly dextrinoid. Stipe vesture 
common, composed of clavate, cylindrical to fusoid or irregular in outline, 
18.4-32 x 8-12 um, hyaline, thin-walled. Clamp connections present. 


oS) 
ie) 


Species of Marasmius (Agaricales: Tricholomataceae) 


Specimen examined: INDONESIA. East Kalimantan, Kayan Mentarang 
National Park, forest surrounding Pa’raye village, 4 Apr 2003, A. Retnowati 
391 (BO, SFSU). 


Distribution: Japan (type), Malaysia, Thailand, Republic of Korea, Papua 
New Guinea, and Indonesia. 


Habit and habitat: Solitary on soil. 


Figure 1. Marasmius cf. purpureostriatus Hongo (A.Retnowati 39/). A. Basidiomes; B. Basidioles; 
C. Cheilocystidia; D. Pileipellis; E. Caulocystidia. 


Notes: M. cf. purpureostriatus is characterized by several peculiar features, 
which include relatively large, greyish magenta pileus, sulcate, distant 
lamellae and absence of pleurocystidia. Basidiospores of the material 
examined are unobserved, but other characters match with description of 
M. purpureostriatus (fide Desjardin and Horak, 1997). As a result, I identify 
the material as M. cf. purpureostriatus. 

M. purpureostriatus 1s allied with M. musisporus from Papua New 
Guinea (Desjardin and Horak, 1997). M. musisporus differs in forming 
smaller basidiome (8-25 mm diam), paller pileus, and the absence of stipe 
vesture (Desjardin and Horak, 1997). 


34 Gard. Bull. Singapore 62 (1) 2010 


2. Marasmius guyanensis Mont. 
Mont., Ann. Sci. Nat. Bot. 4(1): 114. 1854. -Type: French Guyana, Leprieur, 
Dennis 255. (Holo, K). Fig. 2. 


Pileus 2-5 mm diam, convex, with a small dark papilla at the depressed 
center; margin sulcate, straight, crenate; surface dull, dry, smooth, glabrous; 
orange (6-A6). Context thin, white. Lamellae adnate to a collarium, distant 
(7-9 reaching stipe) with no lamellulae, narrow, white to off-white, non- 
marginate. Stipe 5-7 x 0.1 mm, central, terete, pliant, smooth, glabrous, shiny, 
insititious, cylindrical; dark brown to black; no-nodes, stipe arises from dark 
rhizomorphs, black rhizomorphs association. Odor and taste not distinctive. 
Basidiospores (11.2)12-13.6 x 4 um [x = 12.83 + 0.67x 4 +0 um, QO = 2.80- 
3.40, q= 3.21 + 0.17, n = 25 spores per one specimen], elongate-ellipsoid, 
smooth, hyaline, inamyloid, thin-walled. Basidia unobserved. Basidioles 
fusoid, clavate. Cheilocystidia common, composed of Siccus-type broom 
cells; main body 10.4-23.2 x 6.4-9.6 um, cylindrical to clavate, broadly clavate, 
subglobose or irregular in outline; hyaline, thin walled; apical setulae 1.6-3.2 
x 0.8 um, cylindrical to conical, subacute to obtuse or irregular in outline, 
hyaline, thin-walled. Pleurocystidia absent. Pileipellis hymeniform, mottled, 
composed of Siccus-type broom cells; main body 8.8-16 x 5.6-10.4 um, 
cylindrical, clavate, broadly clavate, to subglobose or irregular in outline, 
hyaline, thin-walled; apical setulae 2.4-3.2 x 0.8 um, cylindrical to conical, 
obtuse to acute; hyaline, thin-walled. Stipe tissue monomitic; cortical 
hyphae 3.2-5.6 um diam, parallel, cylindrical, smooth, hyaline; dextrinoid, 
thin-walled; medullary hyphae 2.4-7.2 um diam, cylindrical, dextrinoid, thin- 
walled. Stipe vesture absent. Clamp connections present. 


Specimens examined: INDONESIA. Java, Bogor Botanical Garden, 7 Jan 
1998, D.E. Desjardin 6713 (BO 98-4, SFSU); same location, 8 Jan 1998, D.E. 
Desjardin 6719 (BO 98-19, SFSU);same location, 15 Jan 1998, D. E. Desjardin 
6797 (BO 98-198, SFSU); same location, 12 Jan 1999, A. Retnowati 125 (BO 
99-245, SFSU). East Kalimantan, Kayan Mentarang National Park, forest 
surrounding Pa’raye village, 5 Apr 2003, A. Retnowati 412 (BO). 


Distribution: Indonesia, Martinique, Dominica, and French Guiana. 

Habit and habitat: Gregarious on leaves or arising directly from rhizomorphs. 
Notes: Marasmius guyanensis firstly described by Montagne from French 
Guiana (fide Singer, 1976). It is apparently a widespread species throughout 


tropical Central and South America (Pegler, 1983) and Asia (Corner, 1996). 
This species is easily distingushed by having a small, orange pileus with dark 


Species of Marasmius (Agaricales: Tricholomataceae) 35 


Figure 2. Marasmius guyanensis Mont. (A. Retnowati 412). A. Basidiomes; B. Basidiopores; C. 
Basidioles; D. Cheilocystidia; E. Pileipellis. 


papilla at the center, a presence of collarium, and basidiospores 12-13.6 x 4 
uum and it is abundant in research sites. 


3. Marasmius coklatus var. mentarangensis Retnowati, var. nov. 

Pileus latus, convexus, velutinus. Lamellae adnatae, brunneus marginatae. 
Stipes teres aequalis, minute pruinose, haud insititious. Odor saporque 
nulli. Basidiosporae (5.6) 6.4-7.2 x 3.2-4 um, ellipsoideae, leves, hyalinae, 
inamyloideae, tenui-tunicate. Basidiola 21.6-30.4 x 6.4-7.2 pum, anguste 
clavata. Cheilocystidia cellulae typi Sicci, 11.2-4 x 4-8 um, setulosae, clavate 
vel irregulars, hyalinae; 2-4 setulae ad apicem  6.4-13.6 x 0.8-2.4 um. 
Pleurocystidia nulla. Pileipellis hymeniform, cellulae typi Sicci, 12-36 x 4.6-8 
pum; 2-6 setulae ad apicem, 4-16 x 1.6-3.2 um. Gregarius ad terra. -Holotypus: 
Indonesia. East Kalimantan, Kayan Mentarang National Park, 4 Apr 2003, 
A.Retnowati 390 (BO). Fig. 3. 


Pileus 26-80 mm diam, conic at first, then convex in age, hygrophanous; 
margin incurved, surface strongly wrinkled at the disc, velutinous overall; 
dark brown overall (9-F8). Context white, thick up to 6 mm. Lamellae 
adnate, 24-26, closely attached to stipe, with 3 series of lamellulae, narrow, 
dark brown marginate; pure white. Stipe 50-100 x 9-14 mm, hollow, cylindrical, 
equal, center, non-insititious, smooth, pruinose; light brown. Odor and taste 


36 Gard. Bull. Singapore 62 (1) 2010 


Figure 3. Marasmius coklatus var. mentarangensis Retnowati, var. nov. (A. Retnowati 390, 
Holotype). A. Basidiomes; B. Basidiospores; C. Basidia and Basidioles; D. Cheilocystidia. E. 
Pileipellis; F. Caulocystidia. 


not distinctive. Basidiospores (5.6) 6.4-7.2. x 3.2-4 um (x = 6.46 + 0.51 x 
3,39. + 0:35, O = 1.60- 2.2 5, gq =1.92 = 0:17; mn = 25ispores pen lespecimen): 
ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia 21.6-30.4 x 6.4-7.2 
um, clavate, 4-spored. Basidioles clavate. Cheilocystidia common, composed 
of Siccus-type broom cells, main body 11.2-4 x 4-8 um, clavate, subcylindrical 
to cylindrical or irregular in outline, yellowish brown, thin to thick-walled; 
apical setulae 6.4-13.6 x 0.8-2.4 um, 2-4 setulae, narrowly cylindrical to 
conical or irregular in outline, subacute to acute, hyaline, thin to thick- 
walled. Pleurocystidia absent. Pileipellis hymeniform, mottled, composed of 
Siccus-type broom cells; main body 12-36 x 4.6-8 tm, clavate, subclavate, 
cylindrical to subcylindrical or irregular in outline, yellowish brown, thin to 
thick-walled; apicals setulae 4-16 x 1.6-3.2 um, 2-6 setulae, often branched 
at the apex, narrowly cylindrical to irregular in outline, yellowish brown, 
thick-walled. Stipe tissue monomitic; cortical hyphae and medullary hyphae 
undifferentiated, 6.2-12 um diam, hyaline to weakly yellowish brown, 
thin-walled, dextrinoid. Stipe vesture common, composed of two types of 
caulocystidia: a) Siccus-type broom cells, main body 20-66.4 x 3.2-4.8 um, 
clavate, clavate to broadly clavate, cylindrical or irregular in outline, hyaline, 


Species of Marasmius (Agaricales: Tricholomataceae) 337) 


thin-walled; apical setulae 3.2-6.4 x 1.6 um, not crowded, narrowly cylindrical 
to conical, obtuse to acute, thin to thick-walled up to 0.8 um; b) non-setulae, 
main body 35-36 x 5.6 um, clavate, cylindrical, thin-walled, hyaline. Clamp 
connections present. 


Distribution: Indonesia. 
Habit and habitat: Gregarious on soil. 


Notes: Marasmius coklatus was described from Cibodas Botanical Garden 
by Desjardin, Retnowati and Horak (2000). This javanese materials have a 
dark chocolate brown, velutinous pileus, remote to distant, broad, greyish 
brown lamellae, a brownish orange to brown, pruinose stipe, moderately 
long and broad basidiospores, Siccus-type cheilocystidia and pleurocystidia 
with few (2-5), long (-30 um) apical setulae and numerous pileo-, cheilo-, 
pleuro-, and caulosetae. The material collected from KMNP differs from 
the Javanese material in having shorter basidiospores (6.4-7.2 x 3.2-4) um, 
and the absence of pleurocystidia. These two characters separate the KMNP 
collection as a new variety of M. coklatus. 


4. Marasmius caryote var. pa’rayeensis Retnowati, var. nov. 

Differt a pileo sulcatus, lamellis numerosus, basidiosporis longissimus 
(28)30.4-33.8 x 4.8-5.6 um longis), stipitis caulocystidiis cylindricis deim 
ordinary. Sparsus ad solum. —Holotypus: Indonesia, East Kalimantan, 
Kayan Mentarang National Park, forest along Parinibung, 10 Apr 2003, 
A. Retnowati 460 (BO). Fig. 4. 


Pileus 17-48 mm diam, convex at first, then convex with upturned margin 
in age; hygrophanous, sulcate; margin straight at first, upturned in age; 
surface glabrous, smooth; light brown. Context thin, light brown. Lamellae 
adnate, close (18 reached stipe) with 2-3 series of lamellulae, narrow, non- 
marginate; light brown. Stipe 100-145 x 1.5-2 mm, equal, cylindrical, central, 
non-insititious; smooth, pruinose; light brown to dark brown; white base 
tomentose at the base. Odor and taste not distinctive. Basidiospores (28 )30.4- 
B55 645-50) fon | x— 31.85 += 1.42 x 5.12 + 0.40 pm; O = 5.43-7, g = 6.25 + 
0.51, n = 25 spores per | specimen], cylindrical, smooth, hyaline, inamyloid, 
thin-walled. Basidia 42.4-44 x 10.4 um, 4-spored, clavate. Basidioles clavate. 
Cheilocystidia abundant, 10.4-28 x 8-12.8 um, clavate, broadly clavate to 
pyriform or globose, hyaline to weakly yellow, thin-walled. Pleurocystidia 
absent. Pileipellis hymeniform, not mottled, 12-21.6 x 5.6-9.6 um, clavate 
to broadly clavate, subglobose or globose, hyaline, thin-walled. Stipe tissue 
monomitic; cortical hyphae 7.2-16.8 tm diam, parallel, cylindrical, hyaline, 


38 Gard. Bull. Singapore 62 (1) 2010 


thin- walled,smooth, weakly dextrinoid; medullary hyphae 5.6-10.4 um diam, 
parallel, cylindrical, hyaline to yellowish brown, thin-walled, dextrinoid. 
Stipe vesture common, 16.8-40 x 4.8-7.2 um, cylindrical to clavate or irregular 
shape, hyaline, thin-walled. Clamp connections present. 


Figure 4. Marasmius caryote var. pa’rayeensis Retnowati, var. nov. (A. Retnowati 460, Holotype). 
A. Basiodiomes; B. Basidiospores; C. Basidia and basidioles; D. Cheilocystidia; E. Pileipellis; 
F. Caulocystidia. 


Distribution: Indonesia. 
Habit and habitat: Scattered on soil. 


Notes: Some species of Marasmius, e.g., M. bekolacongoli Beeli (fide Singer, 
1965) and M. titanosporus Reid & Guillarmod (Reid, 1988), are closely 
related to the M. caryote var. pa’rayeensis; however, the Kalimantan material 
differs in having longer basidiospores from M. bekolacongoli (16.5-23(-28) 
x 3.5-5 um), and shorter basidiospores from M. titanosporus (22-36 x 5-7.2 
um). In addition, the presence of stipe vesture in this Marasmius makes it a 
new variety different from the related species. M. caryote variety typical had 
been reported from Krakatau Island by Boedijn (1940). 


Species of Marasmius (Agaricales: Tricholomataceae) 39 


5. Marasmius gypseus Retnowati, sp. nov. 

Pileus 22-41 mm latus, convexus, hygrophanous, crenatus, glabrus, gypseus. 
Lamella adnate, distantes, haud marginatae. Stipes 40-60 x 1.5-3 mm, aequalis, 
glabrus, Haud insititious, brunneus. K-Basidiospores 6.4-7.2 x 3.2-4 um, 
ellipsoideae, leves, hyalinae, inamyloideae. Basidiola clavate, cheilocystidia 
et pleurocystidia nula. Pileipellis hymeniformis, clavate, prelate clavate, 
globosus vel subglobosus, leves, 16-26.4 x 8-18.4 um. Stipitis caulocystidis 
ordinary, 20-24.8 x 6.4-12 um, clavate, globosus vel subglobosus. Cespitosesus 
vel gregarious ad lignum. —Holotypus: Indonesia. East Kalimantan, Kayan 
Mentarang National Park, trail to sub-camp 1 in forest surrounding Pa’raye 
village, 3 Apr 2003, A. Retnowati 371 (BO). Fig. 5. 


Pileus 22-41 mm diam, convex with depressed center, strongly hygrophanous, 
crenate; margin straight, wavy; surface glabrous, wrinkle; white overall. 
Context thin, white. Lamellae adnate, distant with 2 series of lamellulae, 
narrow, non-marginate, white. Stipe 40-60 = 1.5-3 mm, cylindrical, center, 
hollow, equal, mooth, glabrous, non-insititious; brown. Odor and taste not 
distinctive. Basidiospores 6.4-7.2 x 3.2-4 um [xX = 6.78 + 0.41 x 3.33 + 0.30 
um, Q = 1.60-2.25, q = 2.05 + 0.18,n = 25 spores per one specimen], ellipsoid, 
smooth, hyaline, inamyloid, thin-walled. Basidia unobserved. Basidioles 
clavate, fusoid. Cheilocystidia and pleurocystidia absent. Pileipellis 
hymeniform, composed of clavate, broadly clavate, globose to subglobose, 
non-diverticulate, 16-26.4 x 8-18.4 um, hyaline, thin-walled. Stipe tissue 
monomitic; cortical and medullary hyphae indistinguishable, 5.6-12.2 
um diam, parallel, cylindrical, hyaline, smooth, thin-walled, dextrinoid. 
Stipe vesture common, composed of clavate, broadly clavate, globose to 
subglobose, non-diverticulate, scattered, 20-24.8 x 6.4-12 um, hyaline, thin- 
walled. Clamp connections present. 


Distribution: Indonesia. 

Habit and habitat: Caespitose to gregarious on wood. 

Notes: This new Marasmius can be distinguished by having convex with 
depressed center pileus, strongly hygrophanous and distant with 2 series of 


lamellulae, the lacking of cheilocyctidia and pleurocystidia, and basidiospores 
6.4-7.2 x 3.2-4 um. Several members of section G/obulares have a similar 


40 Gard. Bull. Singapore 62 (1) 2010 


Figure 5. Marasmius gypseus Retnowati, sp. nov. (A. Retnowati 371, Holotype). A. Basidomes; B. 
Basidiopores; C. Basidioles; D. Pileipellis; E. Caulocystidia. 


feature with this Marasmius. M. albertianus Singer described from Congo 
has longer basidiospores 8-10 x (3)-4(-5) um, presence of cheilocystidia, 
and is host specific on bamboo (fide Pegler, 1977). M. niveus from French 
Guyana (fide Singer, 1976) also has longer basidiospores 6.5-9 x 2.5-3.5 um, 
and absence of cheilocystidia and stipe vesture. 


Acknowledgment 


The author thanks to Simon Taka Nuhamara (BIOTROP), Danang Anggoro 
(BKSDA Kalimantan Timur), and Deni Wahyudi (Mulawarman University) 
for help during the fieldwork. I am also thankful to Prof. Dr. Mien A. Rifai 
for his useful comments on this paper, and to Indonesian Plant Taxonomy 
Association (PTTI) which facilitated the editors of scientific journals to 
review this paper. This research was funded by World Wide Fund (WWF) 
Indonesia programme to do fieldwork in Kayan Mentarang National Park 
in East Kalimantan. 


Species of Marasmius (Agaricales: Tricholomataceae) 4] 


References 


Boedijn, K.B. 1940. Mycetozoa, fungi, and lichenes of the Krakatau Group. 
Bulletin du Jardin botanique de Buitenzorg III, 16: 398-399. 


Corner, E.J.H. 1996. The agarics genera Marasmius, Chaetocalathus, 
Crinipellis, Heimiomyces, Resupinatus, Xerula, and Xerulina in Malesia. 
Bethefte zur Nova Hedwigia 111: 1-141. 


Desjardin, D.E. and E. Horak. 1997. Marasmius and Gloiocephala in South 
Pacific Region: Papua New Guinea, New Caledonia, and New Zealand 
Taxa. Bibliotheca Mycologica 168: 1-152 . 


Desjardin, D. E.,A. Retnowati and E. Horak. 2000. Agaricales of Indonesia.. 
2.A preliminary monograph of Marasmius from Java and Bali. Sydowia 
52(2): 92-93. 


Hennings, P. 1900. Fungi Monsunenses. Monsunia 1: 15-16; 150-151. 


Kornerup, A. and J.H. Wanscher. 1978. Methuen Handbook of Colour. 3rd. 
Ed. Eyre Methuen, London. 252 pp. 


Overeem, C. van and D. van Overeem-de Haas. 1922. Verzeichnis der in 
Niederlandisch Ost Indien bis dem Jahre 1920 gefundenen Myxomycetes, 
Fungi, und Lichens. Bulletin du Jardin botanique de Buitenzorg III, 4: 88- 
89. 


Léveillé, J.H. 1844. Champignons exotiques. Annales de la Societe des 
Sciences Naturelles (Paris), 3. sér., 2: 167-221. 


Léveillé, J.H. 1846. Descriptions des Champignons de |’Herbier du Muséum 
de Paris. Annales de la Societe des Sciences Naturelles (Paris), 3. sér., 5: 
111-167. 


Moritzi, A. 1845-1846. Verzeichnis der von H. Zollinger in den Jahren 1842- 
1844 auf Java gesammelten Planzen. Solothurn. 144 pp. 


Pegler, D.N. 1977. A preliminary agarics flora of East Africa. Kew Bulletin 
Additional Series. 6: 1-615. 


Pegler, D.N. 1983. Agarics flora of Lesser Antilles. Kew Bulletin Additional 
Series. 9: 195-232. 


42 Gard. Bull. Singapore 62 (1) 2010 


Petch, T. 1947. A revision of Ceylon Marasmii. Transactions of the British 
Mycological Society. 31: 19-44. 


Reid. 1988. Marasmius titanosporus, a new species from the eastern Cape, 
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909. 


Singer, R. 1964. Marasmius congolais recueillis par Mme Goosens-Fontana 
et d’autres collecteurs belges. Bulletin du Jardin Botanique de I Etat. 
[Bruxelles.] 34: 317-388. 


Singer, R. 1965. Monographic Studies on South American Basidiomycetes, 
especially those of the east slope of the Andes and Brazil. 2. The genus 
Marasmius in South America. Sydowia 18: 106-358. 


Singer, R. 1976. Marasmieae (Basidiomycetes-7richolomataceae). Flora 
Neotropica Monograph 17: 1-347. 


Takahashi, H. 1999. Marasmius brunneospermus,a new species of Marasmius 
section Globulares from central Honshu, Japan. Mycoscience 40: 477- 
481. 


Takahashi, H. 2000. Three new species of Marasmius section Sicci from 
eastern Honshu, Japan. Mycoscience 41: 313-321. 


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in den Jahren 1842-1848 gesammelten sowie aus Japan empfangenen 
Pflanzen. Zurich. 160 pp. 


Gardens’ Bulletin Singapore 62 (1): 43-51. 2010 43 


Lectotypification of Elatostema subscabrum H.Schroet. 
(Urticaceae) 


JLT. HADIAH AND BJ.CONN> 


‘Kebun Raya Bogor, Jl. Ir. H. Juanda 13, PO Box 309 Bogor 16122, 
Indonesia 
: Email address: jhadiah@yahoo.com 
- National Herbarium of New South Wales, Mrs Macquaries Road, Sydney 
NSW 2000, Australia 
Email address: bauy.conn@bgsycl.nsw.gov.au 


Abstract 


Descriptions of Elatostema sessile J.R.Forst. & G.Forst and E. subscabrum 
H.Schroet.(Urticaceae) are provided to clarify the morphological differences 
between these two species. The lectotype of E. suwbscabrum H.Schroet. 
(Urticaceae) is here selected. 


Introduction 


Elatostema J.R.Forst. & G.Forst. (Urticaceae) is a very speciose genus of at 
least 300 species occurring throughout the Old World tropics, subtropics and 
subtemperate regions. The general lack of knowledge about the systematics 
and circumscriptions of the species has resulted in the frequent incorrect 
application of plant names because there are no recent publications on 
the systematics of this group. The most recent account of the genus by 
Schréter and Winkler (1936) was based on the relatively few collections 
that were available. Furthermore, this publication only described species 
of subgenera Elatostematoides, Pellionia, and Weddellia (as circumscribed 
by them). Species of Elatostema sensu stricto, that Schroter and Winkler 
(1935) regarded as consisting of 240 species, were not included in their 
publication. 

Recently, progress has been made on our understanding of the 
taxonomy and phylogeny of the Urticaceae (Friis, 1993; Hadiah er al., 
2003, 2008; Sytsma et al., 2002; Wilmot-Dear, 2009) and tribes, in particular 
Elatostemateae (sensu Conn and Hadiah, 2009). Traditionally, the 
taxonomic studies of the genus were based on morphological characteristics 
(Robinson, 1910, 1911; Schréter and Winkler, 1935, 1936; and Weddell, 1854, 
1856, 1857, 1869). A detailed discussion of the morphological characteristics 
used to circumscribe taxa within Elatostema and their usefulness in re- 


44 Gard. Bull. Singapore 62 (1) 2010 


construction of phylogenies are presented in Hadiah and Conn (2009). 
There has been a large volume of work on the taxonomy of the Chinese 
species of Elatostema sensu lato (Wang, 1980a, 1980b; Yang et al., 1995), 
one of the centres of species diversity for the genus. However, little has 
been published on the Malesian species, which are part of another major 
centre of diversity. With relevance to this paper, the above studies provide 
an, albeit incomplete, framework for further systematic study of the genus. 

During studies of the Malaysian species of Urticaceae, material of 
E. sessile has often been confused with E. subscabrum. Two herbarium 
sheets of type material of E. subscabrum were located at the herbarium 
of the Singapore Botanic Gardens (SING). These specimens clarify the 
circumscription of the latter species. 


Elatostema subscabrum H.Schroet. 
Repertorium specierum novarum regni vegetabilis 83: 1 (1935) 20 in obs.; 
Repertorium specierum novarum regni vegetabilis 83:2 (1936) 85. 


Lectotype (here chosen): Malaysia, Pahang, Telom, Nov 1908, H.N. Ridley 
13789 (upper left specimen, SING s.n.); isolectotype (lower right specimen, 
SING s.n.). Other syntype material: H.N. Ridley 13789 (SING67093) (refer 
to discussion of lectotypification, below). 


Terrestrial herb, 0.15-0.2 m_ high, self-supporting (erect/suberect) or 
spreading; internodes developed (elongate, distinct); branched hairs 
lacking; stinging hairs absent; monoecious. Stipules axillary, interpetiolar, 
persistent, free, 1-1.3 mm long. Leaves opposite, appearing alternate (by 
misinterpretation — nanophylls caducous); petiole absent (or < 2 mm long); 
megaphylls with lamina 23-60 mm long, 12-24 mm wide (length to width 
ratio 1.92-2.5), unequal-sided, larger side of lamina ovate to elliptic, smaller 
side elliptic to obovate; surface flat, not rugose, with 4 or 5 vein-pairs; 
venation actinodromous (‘type IVd, semi pinnate nerved’ sensu Schroter 
and Winkler, 1935); basal pair of secondary veins arising from above base 
of primary vein and arising from different points (more than 2 mm apart), 
both directed towards margin (or almost so), joined to next distal secondary 
vein; abaxial surface with cystoliths present on interstices (cystoliths linear), 
with hairs on primary, secondary and tertiary veins; adaxial surface lacking 
cystoliths, glabrous; base oblique, rounded or cuneate; margin toothed, not 
lobed, glabrous; apex short-acuminate; nanophylls not known. Inflorescences 
unisexual; male inflorescences not known; female inflorescence subsessile 
to shortly pedunculate, unbranched, head-like, involucral bracts narrowly 
ovate, sparely hairy to almost glabrous; bracteoles very long, narrowly 
ovate; flowers condensed/crowded, unisexual; actinomorphic (or slightly 


Lh 


Lectoiypification of Elatostema subscabrum 45 


asymetrical): tepals 4. free. unequal, one tepal c. 1mm long with appendage 
1-1.5 mm long: 3 tepals slightly smaller, ca 0.75 mm long without appendage: 
staminodes present. 4, inflexed in bud: ovary straight: style absent: stigma 
oblong, filiform to linear. Achene not enclosed (or only partly so): surface 
ribbed or punctate. 


Lectotypification of Elatostema subscabrum H.Schroet. 

Hilde Schréter described the new species E. subscabrum from material 
collected from Telom, Pahang, Malaysia, with the type cited as Ridley 
13789 (SING) (Schroéter and Winkler, 1936). Two sheets of syntype material 
are held at SING. One herbarium sheet is annotated: *H.N. Ridley 13789. 
Noy, 1908, Telom [all written in Ridley’s hand]. State of Pahang, Malay 
Peninsula [State and region part of pre-printed label — lower right of 
sheet]? (SING67093), with field identification *Elatostema ?acuminatum™ 
[in Ridley’s hand] has three separate samples of the plant: all specimens 
are female and largely in fruit (Fig. 1). The second sheet. °H.N. Ridley 
13789, Nov, 1908, Telom, State of Pahang, Malay Peninsula’ [as annotated 
above — upper left of sheet] (SING), except identified as *E. acuminatum. 
Brongn. by ‘CXF’ [Caetano Xavier Furtado, Singapore Botanic Gardens, 
1923-1952, then 1956-1960] and the label has been stamped “30 MAR, 
1909, presumably the date when the material was accessioned into the 
herbarium (Fig. 2). This second sheet has two female specimens, upper left 
with flowers and fruits, and the lower right specimen largely in fruit. Both 
sheets have been examined by Schréter and annotated as “Elatostema 
subscabrum Hilde Schroter [in her hand], dated *1935° [in her hand]. and 
‘det. Hilde Schréter [pre-printed determinavit slip]. Furthermore, Schréter 
notes that she has not examined male inflorescences (refer protologue), 
only female inflorescence, female flowers and fruit. The upper left specimen 
of the latter sheet (SING) is a good match for the protologue. Therefore, it 
is here selected as the lectotype because it has both flowers and fruits (in 
accordance with Articles 7.11, 8.1-8.3: McNeil er al., 2006). 


Lectotype (chosen by Florence, 1997): J.G.A. Forster [186], Société, Tahiti (P- 
Forst): isolectotype: Forster sn. (BM, K): probably isolectotype: *Dorstenia 
pubescens’ (MW). For further discussion refer Nicolson and Fosberg (2003, 
pp. 677 & 678). 


Distribution: Malaysia; Malay Peninsula, Pahang — only known from type. 
Notes: Collections of E. sessile from Malaysia have frequently been mis- 


identified as E. subscabrum: however, the two species can be distinguished 
by the morphological characters listed in Table 1 (below). 


46 Gard. Bull. Singapore 62 (1) 2010 


: ale ey Atlan fl 
Wy bed Y 


Rass hla 


ee 
Herb. Hort. Bot 
067093 


Singaporensis 


Figure 1. Isolectotypes of Elatostema subscabrum H.Schroet. (Urticaceae) (SING67093). 


Lectotypification of Elatostema subscabrum 


A. neccsrecsc ature, Ssergn. 
both — 


a P 
Gene, AUctrdabntus 
: kL ShcAer 


‘* 


Was. ter Fifide Schroter 


Figure 2. Lectotype (upper left) and isolectotype (lower) of Elatostema subscabrum 
H.Schroet. (Urticaceae) (SING s.n.). 


47 


48 Gard. Bull. Singapore 62 (1) 2010 


Table 1. Diagnostic morphological features distinguishing Elatostema subscabrum from 


E. sessile. 


Character E. subscabrum E. sessile 


Appearing ‘hairy’ because Not appearing ‘hairy’ 
Female of very long, narrowly because of shorter 
inflorescences ovate, tapering, unequal bracteoles not extending 
bracteoles the involucral bracts 


4; unequal size, one of 
which is longer with a long 
appendage 


3; unequal, without 
appendage 


Tepals of female 
flower 


Female flower tepal 


Ree Distinct and readily visible | Minute, difficult to observe 


Elatostema sessile J.R.Forst. & G.Forst. 
Characteres Generum Plantarum 53, n. 2 (1775); ed. 2 (1776) 106. 


Terrestrial herb, 0.3-0.5 m high, self-supporting (erect/suberect); internodes 
developed (elongate, distinct); branched hairs lacking; stinging hairs absent; 
monoecious. Stipules caducous. Leaves opposite, appearing alternate (by 
misinterpretation — nanophylls caducous); sessile (or petiole < 2 mm long); 
megaphylls (45-)50-135(-155) mm long, (21-)22-55(-64) mm wide (length 
to width ratio (2.1-)2.2-2.5), unequal-sided, larger side of lamina ovate to 
elliptic, smaller side as for larger side or to slightly obovate; surface smooth, 
not rugose, with 4 or 5 vein-pairs; venation asymmetric, actinodromous; 
basal pair of secondary veins arising above base of primary vein and arising 
from one point (or less than 2 mm apart), both directed towards apex (or 
almost so), joined to next distal secondary vein; abaxial surface with linear 
cystoliths on interstices and hairs on primary, secondary and tertiary veins; 
adaxial surface with linear cystoliths interstices and on primary, secondary 
and tertiary veins, hairs occasionally present on interstices; base oblique, 
rounded or cuneate; margin toothed, not lobed, sparsely hairy; apex 
acuminate; nanophylls absent. Inflorescences unisexual; male inflorescences 
(based on Weddell, 1869; not known by current authors) sessile or shortly 
pedunculate, depressed-globose, involucral bracts present, broadly 
ovate, hairy on margin; female inflorescence sessile or sometimes shortly 
pedunculate, unbranched, discoid; involucral bracts present, with margin 
hairy, with long appendage; bracteoles linear-spathulate, ciliate; flowers 


Lectotypification of Elatostema subscabrum 49 


condensed/crowded, unisexual: actinomorphic (or slightly asymetrical): 
tepals 3, very minute (appearing absent), unequal, free, appendage absent: 
staminodes 3, inflexed in bud; ovary straight: style absent: stigma oblong. 
filiform to linear. Achene 0.4-0.63 x 0.25-0.35 mm not enclosed (or only 
partly so); surface smooth or ribbed. 


Selected specimens examined: MALAYSIA. Perak: Rotan Segar limestone 
Hill near Tambun, 29 Nov 1960, Allen 4652 (SING): Maxwell’s Hill. 7 Dec 
1965, Shah & Sidek 1146 (SING): Ah Kee Iron Mine, Ipoh, 21 Oct 1958, 
Sinclair 9890 (SING). INDONESIA. Sumatera Utara: Kabupaten Karo, Air 
terjun Sikulikap, Desa Mejuah-juah, 21 Jun 2001 Hadiah, Conn & Ariyanti 
453 (NSW): Jawa Barat: Gunung Gede-Pangrango National Park, track 
to air terjun Cibeureum, 25 Aug 1998, Hadiah 148 (NSW): Bogor, Wana 
Wisata Curug Nangka, Curug Sawer, Warung Loa, 29 Sep 1998, Hadiah 253 
(NSW). 


Distribution: Malaysia. Malay Peninsula: Perak, Perlis, Pahang and Kelantan. 
Indonesia. Sumatera: Sumatera Utara, Sumatera Barat and Jambi; Jawa: 
Jawa Barat, Jawa Tengah and Jawa Timur: Bali. 


Acknowledgements 


This research project was supported by two Singapore Botanic Gardens 
Research Fellowships and one of us (BJC) was also a recipient of a Royal 
Botanic Gardens Sydney Friends’ Science Scholarship. We gratefully 
acknowledge the generous support of these two organisations. 


References 


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Florence, J. 1997. Flore de la Polynesiae francaise. Volume 1. ORSTROM, 
Paris. 


Forster. J.R. and G. Forster. 1775. 53. Elatostema. Characteres generum 
plantarum ... B. White, T. Cadell & P. Elmsly, London. 


Friis, I. 1993. Urticaceae, pp. 612-63. In: Kubitzki, K., J.G. Rohwer, and V. 
Bittrich (eds.). The families and genera of vascular plants II, flowering 
plants — dicotyledones. Magnoliid, hamamelid and caryophyllid families. 
Springer-Verlag, Berlin. 


50 Gard. Bull. Singapore 62 (1) 2010 


Hadiah, J.T. and B.J. Conn. 2009. Usefulness of morphological characters for 
infrageneric classification of Elatostema (Urticaceae). Blumea 54: 181- 
191. 


Hadiah, J.T., B.J. Conn and C.J. Quinn. 2008. Infra-familial phylogeny of 
Urticaceae, using chloroplast sequence data. Australian Systematic 
Botany 21: 375-385. 


Hadiah, J.T., C.J. Quinn and BJ. Conn. 2003. Phylogeny of Elatostema 
(Urticaceae) using chloroplast DNA data. Telopea 10: 235-246. 


McNeill, J., FR. Barrie, H.M. Burdet, V. Demoulin, D.L. Hawksworth, K. 
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and N.J.Turland. 2006. International Code of Botanical Nomenclature 
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Nicolson, D.H. and E.R. Fosberg. 2003. The Forsters and the botany of the 
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Robinson, C.B. 1911. Philippine Urticaceae II. The Philippine Journal of 
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Schroter, H. and H. Winkler. 1935. Monographie der gattung Elatostema 
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an 


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=) 


Gardens’ Bulletin Singapore 62 (1): 53-103. 2010 


Nn 
Oo 


Lejeuneaceae subfamily Ptychanthoideae (Hepaticae) 
in West Java 


I HAERIDA ,S.R.GRADSTEIN AND S. S. TJITROSOEDIRDJO~ 


‘ Herbarium Bogoriense, Botany Division, Research Center for Biology 
Indonesian Institute of Sciences, Cibinong Science Center 
: Jl. Raya Jakarta Bogor Km. 46, Cibinong, Bogor 
Department of Systematic Botany, Institute of Plant Sciences, University of 
_ Gottingen, Untere Karspiile 2, 37073 Gottingen, Germany 
* Department of Biology, Faculty of Science and Mathematics 
Bogor Agriculture University, Bogor, and 
South East Asian Regional Center for Tropical Biology (SEAMEO BIOTROP) 
P.O. Box 116, Bogor, Indonesia 


Abstract 


The subfamily Ptychanthoideae of the family Lejeuneaceae (Hepaticae) 
in West Java is still poorly known. A study of Ptychanthoideae in this 
area reveals the occurrence of 26 species, in 8 genera: Acrolejeunea 
(Spruce) Schiffn. (3 species), Archilejeunea (Spruce) Schiffn. (1 species), 
Lopholejeunea (Spruce) Schiffn. (10 species), Mastigolejeunea (Spruce) 
Schiffn. (3 species), Ptychanthus Nees (1 species), Schiffneriolejeunea Verd. 
(2 species), Spruceanthus Verd. (2 species) and Thysananthus Lindenb. 
(4 species). Mastigolejeunea indica and Thysananthus minor are newly 
reported for Java. The altitudinal and geographical ranges of the species 
of Ptychanthoideae from West Java are presented as well as a key to the 
species and an assessment of their phytogeography. It appears that the 
species are rather widespread; none of the species are endemic to Java or 
western Malesia. The widespread distribution of the species is probably due 
to their dispersal by spores, which may be easily carried by the wind over 
long distances, and by the rather old geological age of the group. 


Introduction 


Lejeuneaceae is a large, mostly tropical family of leafy liverworts, 
containing about 1000 species in 75 genera. According to Gradstein et al. 
(2001), Lejeuneaceae 1s subdivided into two subfamilies, Leyjeuneoideae and 
Ptychanthoideae; the latter group is treated in this paper. 
Ptychantheaeis the only tribe of Ptychanthoideae, while Lejeuneoideae 
consists of three tribes, Brachiolejeuneeae, Lejeuneeae and Cololejeuneeae. 


54 Gard. Bull. Singapore 62 (1) 2010 


The subfamily Ptychanthoideae is characterized by the capsule valves 
spreading widely after dehiscence, elaters usually 72 per capsule, underleaves 
undivided, ventral merophytes 4 or more cells wide, and ocelli absent. In 
Lejeuneoideae the capsule valves are not spreading after dehiscence, elaters 
are 34 or less per capsule, underleaves bifid or undivided, ventral merophytes 
usually 1-4 cells wide, and ocelli sometimes present. 

Ptychanthoideae include fewer genera and species than Lejeuneoideae 
and are generally better known (Gradstein, 1991). They contain about 175 
species in 19 genera worldwide; 14 genera occur in tropical America. Most 
species are xerotolerant epiphytes of forest canopies or rather open, wooded 
vegetation, and usually occur at rather low elevation (Thiers and Gradstein, 
1989; Gradstein et al., 2001). 

Based on several references (Verdoorn, 1933, 1934; Mizutani, 1961; 
Gradstein, 1975, 1991, 1994; Gradstein et al. 2002; Zhu and Gradstein, 2005; 
Gradstein et al., 2005; Hasan and Ariyanti, 2004; Ariyanti and Gradstein, 
2007), there are about 30 species of Ptychanthoideae in Java, in 10 
_genera: Acrolejeunea (Spruce) Schiffn, Archilejeunea (Spruce) Schiffn., 
Caudalejeunea _(Steph.) Schiffn., Dendrolejeunea (Spruce) Lacout., 
Lopholejeunea (Spruce) Schiffn., Mastigolejeunea (Spruce) Schiffn., 
Ptychanthus Nees, Schiffneriolejeunea Verd., Spruceanthus Verd., and 
Thysananthus Lindenb. In addition, there is an old, unconfirmed report of 
the Pacific genus, Phaeolejeunea Mizut., from Java (Zhu and Gradstein, 2005, 
p. 88, originally described as Lopholejeunea latistipula (Schiffn.) Schiffn. var. 
minor Schiffn.). 

Information about the distribution of the species of Ptychanthoideae 
within Java is very scarce. The publications on this subject are mostly from 
the 19° or early 20° century and outdated (e.g., Reinwardt, Blume and 
Nees von Esenbeck, 1824; Nees von Esenbeck, 1830; Sande Lacoste, 1856; 
Schiffner, 1898; Stephani, 1912; Verdoorn, 1933, 1934). 

The aim of this study was to improve our understanding of the diversity 
of Ptychanthoideae in West Java (including West Java Province and Banten 
Province). 


Material and Methods 


The study was based on herbarium specimens of Ptychanthoideae collected 
from West Java and Banten province. A total number of 362 specimens 
deposited in the Herbarium Bogoriense (BO) and Herbarium Biotrop 
(BIOT), and some in the Herbarium of the University of Géttingen, 
Germany (GOET) and Herbarium Bangi, Malaysia (UKMB) kept in BIOT 
as duplicates, were examined. Additional specimens were collected by the 


in 
Nn 


Lejeuneaceae subfamily Prychanthoideae 


first author at several locations in West Java, i.e. Telaga Warna, Ciater, Bogor 
Botanical Garden, Bodogol Education Center and Conservation — Mt. Gede 
Pangrango National Park and Mt. Halimun Salak National Park. 

Morphological characters were studied with a 10 x 4,10 x 10,10 x 20 and 
10 x 40 magnification. Specimens with similar characteristic were grouped 
and identified using recent literature on the species of Ptychanthoideae, 
such as Thiers and Gradstein (1989), Gradstein er al. (2001), Gradstein et 
al. (2002) and Zhu and Gradstein (2005). Descriptions and illustrations of 
each species were prepared and an identification key to the species was 
developed. 


Taxonomic treatment 


Key to the species of West Javan Ptychanthoideae 


1. Branching always Frullania type.......................18. Ptychanthus striatus 
Betseancnins Mejeuned OF Friullania ty Pe......:......0.....0aessscssecssssnecsecesese «iene oe: 2 
2. Branching Frullania and Lejeunea type (Schiffneriolejeunea)................3 
PEM AME MEM ILCICUIMCE-LYPC ONY. 2. 55.2.5. 25 058. tects coe owe ee cceseecteesstoccetens - 


3. Lobule forming a distinct sac at the base and flattened above with 2 large 
teeth. Teeth erect, not pointing outwards towards leaf apex. Leaves 
squarrose when moist.. 20. Schiffneriolejeunea tumida var. haskarliana 

3. Lobule not forming a distinct sac at the base, free margin of the lobule 
plane and clearly visible, with 2 teeth. Teeth almost equal in size and 
pointing outwards towards leaf apex. Leaves not squarrose when moist 
Pe BSS s sheds Sn toca avaiees 19. Schiffneriolejeunea pulopenangensis 


4. Lobes with a distinct vitta ...........................25. Thysananthus retusus 
2 DE EC ee ae eee 5 
Pee cls Orthe LODE With COLMAtE ITIDONES.... ......... ..eerecseseerodeececeeeoeeroveeO 
Pecusorte lobe not with cordate trigONes. ...... 15.20.00. sce0e soe eeeeseeeees 14 


6. Leaves and underleaves entire. Dorsal epidermis cells larger than inner 
DEE FELD 2 2S See ee nee een erry aera eee | 
6. Leaves and underleaves usually toothed. Dorsal epidermis cells not larger 
pam eECe Stem CEUS( PH YSAVIGIINUS) - .-< <cc-ncetwennnsenconanracedvewsieenscsens 12 


Lee 
7 


Gard. Bull. Singapore 62 (1) 2010 


. Perianth 3 keels, with 1-2 innovations (Mastigolejeuned).......ccccccescceneen 8 
. Perianth 8-10 keels, innovation lacking (Acrolejeunea)...................... 10 
; Leaf lobule with several large; triangular tecthis2eee eee 

Fe er Pore eet eet eet Mee A Pete 5A 16. Mastigolejeunea indica 

Leaf lobule with one tooth). #c.cge sec: aneks ce eee eee eee eee eee 9 
. Lobule tooth 3-5 cells long, sharp, curved....... 17. Mastigolejeunea virens 
» Lobule tooth Icell long; blumtynot cukved irc eee eee ee 


cabal steve oth Se Sah sd Rupees eetes Meta. cte eee meee 15. Mastigolejeunea auriculata 


. Lobule semicircular in outline, with 4-10 teeth, all or at least some teeth 


23 Cells 1ONGwee eee ae cde eee ee 2. Acrolejeunea fertilis 


-Lobule not semicircular, withi2-siteethi 4.2: ea 


. Lobule narrow rectangular, with 2 teeth ......... 1. Acrolejeunea arcuata 
. Lobule ovate-rectangular, with 3 teeth ......3. Acrolejeunea pycnoclada 


. Plant very small, less than 1 mm wide .............. 24. Thysananthus minor 
/ Plant more than fmm wide .2.4.<2.52..-04 eee eee ee 13 


. Upper part of leaf asymmetrical, turned to the ventral side (“ventrad”’), 


leat apex recive secre. eee eee 23. Thysananthus convolutus 


. Upper part of leaf symmetrical, not turned to the ventral side, leaf apex 


planeionmecuny edie a. eee 26. Thysananthus spathulistipus 


. Stems robust, ventral merophyte 8-12 cells wide (Spruceanthus)........15 
. stems' thinner, ventral metrophyte 4-6 cells widen ce. -ee eee ee 16 


» Keatapexishanply;actitew:--.-ee.s ee 22. Spruceanthus semirepandus 
; beat apexsrounded (oo btusem=e = 21. Spruceanthus polymorphus 


. Stem epidermis cells not larger than the inner cells. Plants light green. 


Perianth keels smooth or weakly toothed. Innovations present.............. 
sett osieudeuberubedaenies eeeanee eee ete ce eee eee 4. Archilejeunea planiuscula 


. Stem epidermis cells larger than the inner cells. Plants blackish-green, 


rarely brown. Perianth keels distinctly toothed (Lopholejeunea). 
Innovations lackim pst: -sec.eces-s eeneree ete eee 17 


Leaf lobule connected to the leaf lobe by only one single cell............18 
Leaf lobule connected to the leaf lobe across several cells.................21 


Lejeuneaceae subfamily Ptychanthoideae 57 


22: 


24. 


24. 


25. 


. Leaf lobe acute .....00.0. 0.0). Lopholejeunea applanata 
BEcaAnOuE LOUnGCEd OF ODIUSE.....25:22.2-..edeeten se 0b edoncse ce cWecee te dedeeee 19 
. Lobule deeply constricted in the middle, free margin strongly involute 


ee ec asin oe pds ene cove sue ncvtendcassaves sade 7. Lopholejeunea herzogiana 


. Lobule not deeply constricted in the middle, free margin not involute.....20 


. Underleaves distant to contiguous, orbicular. Female bracteole entire 


Seno ehecnp scan Sancnstbesbnaitcsuhonssonion 8. Lopholejeunea nigricans 


. Underleaves imbricate, wider than long. Female bracteole toothed........ 


-c sserseceeec oe eee re 6. Lopholejeunea eulopha 


. Underleaves very large, 6-8 x as wide as the stem ..... ey 
meacericaves smaller 4-6 x as wide as the Stem ............2......:.s000000.- 23 


. Plants pale brown in the dried condition. Leaf lobules small, 1/5-1/4 of 


feat leapth Female bracteole toothed. Dioicous ..........2.........<.ccessessecessssees 
Se cles Soha dnc vas onedncvnsnasneve’ 13. Lopholejeunea wiltensii 
Plants blackish or dark brown to blackish in the dried condition. Leaf 
lobules larger, 1/3-2/5 of leaf length. Female bracteole usually entire. 
8 ICDS TE ae Be ee 14. Lopholejeunea zollingeri 


3. Plants minute, less than 1 mm wide............. 10. Lopholejeunea horticola 
miaaes medium sized, more than 1 mm wide... ......-..-...2..0.0..se0ssseee8 24 


Margins of underleaves and leaves recurved. Underleaf large, more 
eee ee ROD EMCH PLN: «ci. 25. c.cccveasesessceeeseco = 28 11. Lopholejeunea recurvata 
Margins of underleaves and leaves plane. Underleaf less than % x 
C2 DE EOD ae scence sp ere ee oe Ay 2 


. Female bracteole entire. Female bract lobule very small or lacking 


ses ne neecececiese Coane Oe 12. Lopholejeunea subfusca 
Female bracteole crenate. Female bract lobule large... ee 
se saghiss nda se ec Ocae 9. Lopholejeunea ceylanica 


Species descriptions 


1. Acrolejeunea arcuata (Nees) Grolle & Gradst., J. Hattori Bot. Lab. 38: 332 
(1974). 

Jungermannia arcuata Nees, Enum. Pl. Crypt. Javae 1:38 (1830). Ptychocoleus 
arcuatus (Nees) Trevis., Mem. Reale Ist. Lomb. Sci. Mat. Nat., Ser. 3, 4: 405 


58 Gard. Bull. Singapore 62 (1) 2010 


(1877). -Type: Indonesia. Java: Lebak Mts., Reinwardt s.n. (holotype, STR; 
isotypes, G, NY, W) — cf. Gradstein et al. (2002). 


Figure 1. Acrolejeunea arcuata (Nees) Grolle & Gradst. Shoot (a); leaf-lobe (b); lobule (c); cells of 
midleaf (d); underleaf (e). Drawn from Kornochalert 1408, BIOT. 


Plants up to 2 cm long, about 0.9 mm wide, dark brown to reddish brown in 
dried condition. Branching Lejeunea-type. Stem diameter 95-108 um. Leaves 
closely imbricate, obliquely spreading. Lobe ovate-orbicular to obovate, 520- 
650 um long, 410-520 um wide, margin entire, apex subobtuse to sometimes 
rounded; cells of lobe thin-walled, hyaline, somewhat elongate, trigones + 
cordate, intermediate thickening scarce; marginal cells 5-11 x 6-11 um, mid- 
leaf cells 19-29 x 11-26 um, basal cells 24-30 x 10-13 um; oil bodies not seen. 
Lobule narrow rectangular, 300-400 um long, 145-160 um wide, apex truncate 
with 2 teeth, each tooth consisting of 2-3 cells, 2-3 cells long. Underleaves 
closely imbricate, subtriangular to suborbicular, 260-315 tm long, 295-435 
um wide, margins entire, apex truncate, base slightly auriculate, insertion 
line shallowly curved. Generative structures not seen. 


Distribution: Java, Sumatra, Borneo, Philippines, Papua, Peninsular Malaysia. 


Specimen examined: INDONESIA. West Java, Mt. Patuha, on bark of tree 
along the road to crater, 2000 m, Kornochalert 1408 (BIOT!). 


Notes: Diagnostic characters of Acrolejeunea arcuata are: 1) lobules 2(-3) 
times longer than wide, with 2 teeth, 2) lobe ovate-orbicular with entire 
margins, 3) cordate trigones. This species was found at ca 2000 m and occurs 
only at higher elevations in the mountains (Gradstein, 1975). It is readily 


in 
\O 


Lejeuneaceae subfamily Prychanthoideae 


distinguished from other species of Acrolejeunea by the very long and 
narrow lobule with 2 teeth and the often reddish-brown color. 


2. Acrolejeunea fertilis (Reinw. et al.) Schiffn., in Engler and Prantl, Nat. 
Pflanzenfam. 1, 3: 128 (1893); Jungermannia fertilis Reinw. et al., Nova Acta 
Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 211 (1824): Ptychocoleus 
fertilis (Reinw. et al.) Trevis., Mem. Reale Ist. Lomb.Sci. Mat. Nat., Ser. 3, 4: 
405 (1877). —Type: Indonesia. Java, Lebak Mts., Reinwardt s.n. (holotype, 
STR; isotypes, G, NY, W) — cf. Gradstein et al. (2002). 


Figure 2. Acrolejeunea fertilis (Reinw. et al.) Schiffn. Shoot (a): leaf-lobe (b); underleaf (c): cells of 
midleaf (d); cells of basal part of the leaf (e); Drawn from van Borssum Waalkes 549, BO. 


Plants up to | cm long, about 0.8 mm wide, dark green to dark brown in 
dried condition. Branching Lejeunea-type. Stem diameter 100-124 um. 
Leaves closely imbricate, obliquely spreading, squarrose. Lobe ovate- 
orbicular to subrectangular, 600-825 um long, 500-660 um wide, margin 
entire, apex truncate to sometimes rounded; cells of lobe thin-walled, light- 
yellow, somewhat elongate, trigones cordate, intermediate thickening scarce; 
marginal cells 7-8 x 7-10 um, mid-leaf cells 18-23 x 12-20 um, basal cells 17-30 
x 13-17 um; oil bodies not seen. Lobule semicircular, 200-290 um long, 220- 
250 um wide, apex obliquely truncate with 4-9 teeth, each tooth consisting of 
2-3 cells, 1-2 cells long. Underleaves closely imbricate, broadly orbicular, 300- 
350 um long, 310-570 um wide, margins entire, apex truncate, base cuneate, 
insertion line shallowly curved. Generative structures not seen. 


Specimens examined: INDONESIA. Banten, Pulau Panaitan, van Borssum 
Waalkes 549 (BO!). West Java, Depok, Kampus Universitas Indonesia, 60- 
80 m, Afiatri Putika 80 (BO!). 


60 Gard. Bull. Singapore 62 (1) 2010 


Distribution: Java,Sumatra, Borneo, Philippines, Moluccas, Papua, Peninsular 
Malaysia, Indochina, India, Sri Lanka. 


Notes: Diagnostic characters of Acrolejeunea fertilis are: 1) lobule 
semicircular with 4-9 teeth, 2) lobe ovate-orbicular with entire margins, 3) 
trigones cordate. The species occurs in the lowlands, at ca 60-80 m, and is 
distinguished from other Javanese members of the genus Acrolejeunea by 
the squarrose leaves and the semicircular lobule with 4-9 teeth. 


3. Acrolejeunea pycnoclada (Taylor) Schiffn., in Engler and Prantl, Nat. 
Pflanzenfam. 1, 3: 128 (1893); Ptychanthus pycnocladus Taylor, London J. 
Bot. 5: 385 (1846); Ptychocoleus pycnocladus (Taylor) Steph., Sp. Hepat. 5: 
52 (1912). -Type: Peninsular Malaysia, without loc., Cantor s.n. (holotype, 
FH; isotypes, G, MANCH, NY, S, U, W) — cf. Gradstein et al. (2002). 


Figure 3. Acrolejeunea pycnoclada; (Taylor) Schiffn. Shoot (a); leaf lobe (b); leaf lobule (c): 
underleaf (d); leaf cells (e); female bract (f); female bracteole (g); perianth (h). Drawn from Haerida 
1484, BO. 


Autoicous. Plants up to 2 cm long, 0.5-0.7 mm wide, 1-1.5 mm wide in 
wet condition; dull green when fresh, greenish-brown in dried condition. 
Branching Lejeunea-type. Stem diameter 100-145 um; cross-section of the 
stem with epidermis cells larger than medulla cells Leaves imbricate, widely 
spreading, convolute when dry. Lobe ovate, 680- 1050 um long, 370-850 um 
wide, margin entire, apex rounded; cells of lobe thin-walled, hyaline, in mid- 


Lejeuneaceae subfamily Ptychanthoideae 61 


leaf elongate, trigones cordate, intermediate thickening frequent; marginal 
cells 7-16 x 6-13 um, mid-leaf cells 17-31 x 11-25 um, basal cells 24-52 x 
11- 31 um; oil bodies (7-)8-20 per cell, homogeneous, hyaline, globose to 
ellipsoidal. Lobule ovate to rectangular, large, 365-680 um long, 170-310 um 
wide, inflated, apex obliquely truncate with 2 conspicuous teeth, first tooth 
consisting of 3-4 cells, second tooth consisting of 2-3 cells, an additional 
small tooth present at the distal end of the free margin beyond the first 
tooth. Underleaves distant to contiguous, obovate-orbicular to wider than 
long, 270-480 um long, 350-700 um wide, margin entire, apex rounded to 
almost truncate, base cuneate, insertion line shallowly curved. Androecia 
intercalary on branches, bracts in 3-11 pairs, very similar to leaves but 
lobules slightly larger, 400-420 um long, 180-250 um wide, bract lobule 1/2- 
2/3 of the bract lobe, epistatic, bracteoles similar to underleaves in size and 
shape. Gynoecia on short or long branches, innovations lacking, bracts and 
bracteoles in 2 pairs, bract oblong to broadly oblong 720-1070 um long, 
270-500 um wide, conspicuously incurved in the upper part, margin entire, 
apex obtuse to orbicular, sinus up to 1/3 of lobe length; bracteole oblong to 
broadly oblong 890-1010 um long, 520-660 um wide, margin entire to slightly 
undulate, apex truncate. Perianth immersed, obovate, 975 um long, 430 um 
wide, inflated, with + 10 inflated keels, margins entire. Sporophytes and 
asexual reproduction not seen. 


Distribution: Java, Sumatra, Borneo, Moluccas, Philippines, Papua New 
Guinea, Pacific, Peninsular Malaysia, Thailand, India, Sri Lanka, tropical 
Africa. 


Specimens examined: INDONESIA. West Java, G. Gede, above Sukabumi, 
1500-1900 m, Verdoorn 46b (BO!); Ciater, Subang, Sari Ater, ca 1500 m, 
Haerida 1480, 1481, 1484, 1486, 1487 (BO!). 


Notes: Diagnostic characters of Acrolejeunea pycnoclada are: 1) lobule with 
2 conspicuous teeth and an additional, third small tooth present at the distal 
end of the free margin, beyond the first tooth, 2) trigones cordate, 3) female 
bracts conspicuously incurved in the upper part, covering the immersed 
perianth, 4) perianth with 10 keels. This species was found at ca 1500-1900 
m but it has also been recorded at lower elevations, from sealevel upwards. 
It differs from other members of the genus Acrolejeunea by the lobule 
with 2 conspicuous teeth and a 3° small tooth at the very end of the free 
margin, near the junction with the ventral margin of the leaf lobe, and by the 
conspicuously incurved female bracts. 


4. Archilejeunea planiuscula (Mitt.) Steph., Sp. Hepat. 4: 731 (1911); 
Lejeunea planiuscula Mitt., J. Proc. Lin.. Soc. 5: 111 (1861). -Type: Burma. 


62 Gard. Bull. Singapore 62 (1) 2010 


Rangoon, McClelland s.n. (holotype, NY) — cf. Thiers and Gradstein (1989). 
- Archilejeunea caramuensis Steph., Hedwigia 34: 59 (1895). —Type: 
Philippines.Caramuan, 1884-85, Micholitz 70 (G) — cf. Verdoorn (1934). 

- Archilejeunea falcata Steph., Hedwigia 34: 60 (1895). —-Type: Papua New 
Guinea. Madang Prov.: Stephansort, 1888, Kdrnbach s.n. (G) — cf. Verdoorn 
(1934). 

- Mastigolejeunea paradoxaVerd.,Nova Guinea 18:5 (1935).—Type: Indonesia. 
West Irian, Jayawijaya Prov., Prauwenbivak, Lam 1182a (holotype, FH) — cf. 
Gradstein et al. (2002). 


Figure 4. Archilejeunea planiuscula (Mitt.) Steph. Shoot (a); leaf lobule (b); underleaf (c); cells of 
leaf margin (d); cells of midleaf (e). Drawn from Haerida 1489, BO. 


Plants up to 3 cm long, 0.8-2 mm wide; light green when fresh, darker green 
in dried condition. Branching Lejewnea-type. Stem diameter 90-135 um; 
epidermis cells not large than inner cells; ventral merophyte 4-6 cells wide. 
Leaves contiguous, widely spreading. Lobe orbicular to oblong, 500-900 
um long, 370-700 um wide, margin entire, apex rounded; cells of lobe thin- 
walled, hyaline, isodiametrical hexagonal, trigones triangular, never cordate, 
intermediate thickening scarce; marginal cells 3.5-7.5 x 5-10 tm, mid-leaf 
cells 17-24 x 14-18 um, basal cells 22-37 x 13-17 um; oil bodies not seen. 
Lobule ovate, 240-340 um long, 130-200 um wide, apex obliquely truncate 
with 1-2 teeth, first tooth with 2 cells long, second tooth small, 1 cell long, 
sometimes lacking. Underleaves distant to contiguous, obdeltoid, 280-420 
um long, 260-460 um wide, margin entire, apex truncate, insertion line almost 
straight. Generative structures not seen [innovations present; perianth with 
5 smooth or weakly toothed keels]. 


Lejeuneaceae subfamily Prychanthoideae 63 


Specimens examined: INDONESIA. West Java, Bogor Bot. Garden, 
Verdoorn 12a, 12b, 12c, 12e, 131, 134, 232, Meijer B55a, Bola, B49, B76, 
B3664, B27a, B56, BS4a (BO!); Cibodas Bot. Garden, 1450 m, Schiffner 
233, Haerida 813 (BO!); Ujung Kulon Nat. Park, Dewi Dw922 (BO!). 


Distribution: Java, Peninsular Malaysia, Papua New Guinea, Philippines, 
Solomon Is., Pacific, Australia, Indochina, India, Sri Lanka. 


Notes: Diagnostic characters of Archilejeunea planiuscula are: 1) underleaves 
obdeltoid, 2) trigones triangular, never cordate, 3) plant light green in color 
when fresh. This species was found at ca 200-1450 m. It is readily distinguished 
from other members of the subfamily by the rather light green color and 
flat appearance of the plants, lobules with 1-2 teeth, isodiametrical leaf cells 
with small, simple trigones, obdeltoid underleaves and thin stems (ventral 
merophyte 4-6 cells wide) with epidermis cells not larger than inner cells. 


5. Lopholejeunea applanata (Reinw. et al.) Schiffn., in Engler and Prantl, 
Nat. Pflanzenfam. 1, 3: 129 (1893); Jungermannia applanata Reinw. et al. 
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 210 (1824): 
Phragmicoma applanata (Reinw. et al.) Nees, Naturgesch. Eur. Leberm. 
3: 248 (1838); Lejeunea applanata (Reinw. et al.) Nees, in Gottsche et al., 
Syn. Hepat.: 314 (1845): Symbiezidium applanatum (Reinw. et al.) Trevis., 
Mem. Reale Ist. Lombardo Sci., Cl. Sci. Mat., Ser. 3, 4: 403 (1877). —Type: 
Indonesia. Java, collector unknown (holotype, STR: isotypes, G, S, W) — cf. 
Zhu and Gradstein (2005). 

- Lopholejeunea fleischeri Steph., Sp. Hepat. 5: 79. 1912. —Type: Indonesia. 
Java, Tjibodas, Apr 1900, M. Fleischer s.n. (holotype, G) — cf. Zhu and 
Gradstein (2005). 


Plants up to 5 cm long, 1-1.5 mm wide; dark brown in the dried condition. 
Branching Lejeunea-type. Stem diameter 120-200 um Leaves imbricate, 
widely spreading. Lobe ovate, 800-1200 um long, 500-800 um wide, margin 
entire sometimes crenulate towards the apex, apex acute to acuminate: cells 
of lobe thick-walled, yellow, rectangular to isodiametric, trigones triangular, 
intermediate thickening frequent; marginal cells 15-21 x 12-15 um, mid-leaf 
cells 26-30 x 20-24 um, basal cells 30-40 x 20-31 um: oil bodies not seen. 
Lobule ovate, 230-320 um long, 170-220 um wide, inflated, apex truncate 
with 1 small tooth, connected to the leaf lobe across only one single cell. 
Underleaves large, imbricate, broadly orbicular, 300-400 um long, 400- 
900 um wide, margin entire, apex truncate, insertion line deeply curved. 
Androecia intercalary on branches, bracts in 3-12 pairs, ovate with obtuse 
apex, 200-300 um long, 150-220 um wide, bract lobule about % of the lobe 


64 Gard. Bull. Singapore 62 (1) 2010 


length; bracteoles similar to underleaves in size and shape. Gynoecia not 
seen. 


20 Um 


Figure 5. Lopholejeunea applanata (Reinw. et al.) Schiffn. Shoot (a); underleaf (b); leaf lobule 
(c); leaf lobe (d); cells of midleaf (e). Drawn from Meijer 3696, BO. 


Distribution: Java, Sumatra, Borneo, Philippines, Peninsular Malaysia, Papua 
New Guinea, Pacific, Indochina, Sri Lanka, India. 


Specimens examined: INDONESIA. West Java, Cibodas Bot. Garden, 
Meijer B3696 (BO!); above Cibodas, Cibeureum, Hasskarl s.n. (BO!); G. 
Pangrango, “bij Tjisaroea” ca 1200 m, Verdoorn 44b (BO!); G. Patuha, Lake 
Situ Patengan, ca 1500 m, Kornochalert 1413 (BIOT!). 


Notes: Diagnostic characters of Lopholejeunea applanata are: 1) leaf lobes 
acute to acuminate at apex, 2) lobule apex connected to the lobe across only 
one single cell. This species grows at ca 1200-1500 m. It differs from other 
Javanese members of the genus Lopholejeunea by the acute to acuminate 
leaf apex and large, imbricate underleaves. 


6. Lopholejeunea eulopha (Taylor) Schiffn., in Engler and Prantl, Nat. 
Pflanzenfam. 1,3: 129 (1893); Lejeunea eulopha Taylor, London J. Bot. 5: 391 
(1846); Phragmicoma eulopha (Taylor) Mitt.,in Seeman, FI. Vit. 413 (1873); 
Symbiezidium eulophum (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci., 
Cl. Sci. Mat., Ser. 3, 4: 403 (1877). —Type: Pacific islands: locality unknown, 


in 


Lejeuneaceae subfamily Ptychanthoideae 6 


Nightingale s.n. (holotype, FH; isotypes, FH. NY) — cf. Zhu and Gradstein 
(2005). 

- Lopholejeunea nymannii Steph., Sp. Hepat. 5: 84. 1912. -Type: Indonesia. 
Java, G. Salak, E. Nyman s.n. (holotype, G;isotype, W) —cf. Zhu and Gradstein 
2005. 


a 
fi 
a Se 


Figure 6. Lopholejeunea eulopha (Taylor) Schiffn. Shoot (a): leaf lobule (b); leaf lobe (c): cells of 
midleaf (d). Drawn from Haerida 710, BO. 


Dioicous, sometimes autoicous. Plants up to 3 cm long, 1-1.5 mm wide, 
brownish green to dark brown in dried condition. Branching Lejeuwnea-type. 
Stem diameter 130-190 um. Leaves imbricate, widely spreading. Lobe oblong- 
ovate, 430-1090 um long, 340-900 um wide, margin entire, apex obtuse; cells 
of lobe thin-walled, pale brown, hexagonal to nearly isodiametric, trigones 
triangular, intermediate thickening frequent; marginal cells 7-15 x 10-17 um, 
mid-leaf cells 14-31 x 10-21 um, basal cells 22-40 x 18-24 um; oil bodies 
(11-)18-20 per cell, homogeneous, hyaline, ellipsoidal. Lobule ovate, 120-270 
um long, 110-200 um wide, inflated, apex with | triangular tooth consisting 
of 3-5 cells, connected to the leaf lobe by only one single cell. Underleaves 
imbricate, large, reniform, 300-500 um long, 410-1070 um wide, margin entire, 
apex rounded, insertion line deeply curved. Generative structures not seen. 


Specimens examined: INDONESIA. West Java, Bogor Bot. Garden, Meijer 
94b, Schiffner 245, van Steenis s.n. (BO!); Bogor, without locality, Verdoorn 
s.n. (BO!); Cibodas Bot. Garden, 1400 m. Ariyanti 47] (BIOT!); G. Halimun 
Nat. Park, Haerida 710 (BO!); G. Guntur, “bij Kawah Kamoedjan” 1500- 
1700 m.alt., Verdoorn 50h, 50d (BO!); G. Megamendung, Verdoorn 68 (BO!): 


66 Gard. Bull. Singapore 62 (1) 2010 


Kota Batu, Schiffner s.n. (BO!): Kampung Jember, Geger Bentang ca 1350 
m, Neervoort 3087 (BO!): Ujung Kulon Nat. Park, Dewi Dw922d (BO!); G. 
Patuha, Lake Situ Patengan ca 1500 m, Kornochalert 1418, Dian Apriana & 
Afiatri Putrika 25 (BIOT!). 


Distribution: Java, Sumatra, Borneo, Bali, Sulawesi, Moluccas, Philippines, 
West Irian, Papua New Guinea, Solomon Is., Pacific, Australia, Indochina, 
India, Sri Lanka, China, Japan, tropical Africa, tropical America. 


Notes: Diagnostic characters of Lopholejeunea eulopha are: 1) lobule ovate, 
with | triangular tooth consisting of 3-5 cells,2) lobule apex connected to the 
leaf lobe across only one cell, 3) large, reniform underleaves. This common 
species grows at ca 200-1700 m. It resembles L. applanata by the very large, 
imbricate underleaves but differs by the rounded leaf apex. 


7. Lopholejeunea herzogiana Verd., Rec. Trav. Bot. Neerl. 30: 217 (1933). - 
- Type: Indonesia. Java, Cibodas, ‘ad arborum truncos,” 1420 m, 1894, Schiffner 
s.n. (FH, lectotype designated by Zhu and Gradstein, 2005). 
— Lopholejeunea pullei Verd., Nova Guinea 18: 4 (1935). —Type: Indonesia. 
Irian Jaya, Hellwig Mts., “an Gymnospermen”, 2600 m, 10 Jan 1913, Pulle 
s.n. (holotype, FH: isotype, U) — cf. Zhu and Gradstein 2005. 


Figure 7. Lopholejeunea herzogiana Verd. Shoot (a): leaf lobule (b); female bract (c): female 
bracteole (d): cells of midleaf (e). Drawn from Verdoorn 64a, BO. 


Lejeuneaceae subfamily Prychanthoideae 67 


Plants robust, up to 4 cm long, 1-2.5 mm wide: blackish brown in the dried 
condition. Branching Lejeuwnea-type. Stem diameter 130-220 um Leaves 
imbricate, widely spreading. Lobe obovate-triangular to broadly orbicular, 
650-1050 um long, 660-1420 um wide, margin entire, apex obtuse to rounded, 
sometimes recurved; cells of lobe thick-walled, brownish-orange, rectangular 
to hexagonal, trigones triangular, intermediate thickening frequent; marginal 
cells 12-18 x 12-17 um, mid-leaf cells 21-26 x 20-30 um, basal cells 24-50 x 
20-30 um; oil bodies not seen. Lobule ovate, large, 300-540 um long, 220-310 
um wide, inflated, free margin strongly involuted and with a conspicuous 
constriction in the middle, connected to the leaf lobe across only one single 
cell. Underleaves closely imbricate, reniform to transversely rectangular, 
400-520 um long, 600-1310 um wide, margin entire, apex truncate, insertion 
line deeply curved. Androecia not seen. Gynoecia on short or long branches, 
innovations lacking, bracts and bracteoles in 1-2 pairs, bract ovate to nearly 
rectangular, 2-3 mm long, 1.5-2 mm wide, margin toothed, apex truncate, 
sinus up to 1/2 of lobe length, bract lobule ovate, margin ciliate-dentate, 
apex truncate to nearly acute; bracteole oblong 1.3-2.2 mm long, 0.5-1.4 
mm wide, margin toothed, apex rounded. Perianth immersed, obovate, 550 
um long, 530 um wide, inflated, with 5-7 keels, margins widely winged and 
toothed. Sporophytes and asexual reproduction not seen. 


Specimens examined: INDONESIA. West Java, Telaga Warna, Puncak Pass, 
ca 1500 m, Verdoorn 64a (BO!): Cibodas Bot. Garden, Meijer B3800b (BO!): 
G. Pangerango, “bij Tjisaroea”, ca 1200 m, Verdoorn 44c (BO!). 


Distribution: Java, Peninsular Malaysia, West Irian, Papua New Guinea, 
New Caledonia. 


Notes: Diagnostic characters of Lopholejeunea herzogiana are: 1) strongly 
winged and toothed obovate perianth, 2) lobule conspicuously constricted 
in the middle and with strongly involuted free margin, 3) lobule apex 
connected to the leaf lobe across only one single cell, 4) large, + reniform 
underleaf with deeply curved insertion line. This rare species was found at 
ca 1200-1500 m. and usually grows in the mountains at higher elevation. It 
is readily separated from other species of the genus by the peculiar lobule 
with deep constriction in the middle. 


8. Lopholejeunea nigricans (Lindenb.) Schiffn., Consp. Hepat. Arch. Ind. 
293 (1898); Lejeunea nigricans Lindenb., in Gottsche et al., Syn. Hepat. 
316 (1845): Symbiezidium nigricans (Lindenberg) Trevis., Mem. Reale Ist. 
Lombardo Sci., Cl. Sci. Mat., Ser. 3, 4: 403 (1877). —Type: Indonesia. Java, 
without locality, collector unknown (holotype, W: isotypes. G, S) — cf. Zhu 
and Gradstein (2005). 


68 Gard. Bull. Singapore 62 (1) 2010 


— Lejeunea intermedia Lindenb., in Gottsche et al., Syn. Hepat. 316 (1845); 
Symbiezidium intermedium (Lindenberg) Trevis. Mem. Reale Ist. 
Lombardo Sci., Cl. Sci. Mat., Ser. 3, 4: 403 (1877); Lopholejeunea intermedia 
(Lindenberg) Steph., Sp. Hepat. 5:77 (1912). Type: Indonesia. Java, without 
locality, collector unkown (holotype, W) — cf. Zhu and Gradstein 2005. 

— Lejeunea javanica Nees, in Gottsche et al., Syn. Hepat. 320 (1845); 
Symbiezidium javanicum (Nees) Trevis., Mem. Reale Ist. Lombardo Sci., Cl. 
Sci. Mat., Ser. 3, 4: 403 (1877); Lopholejeunea javanica (Nees) Schiffn., in 
Engler and Prantl, Nat. Pflanzenfam. 1,3: 129 (1893). —Type: Indonesia. Java, 
without locality, collector unknown (holotype, W; isotypes, G, S) — cf. Zhu 
and Gradstein (2005). 


Figure 8. Lopholejeunea nigricans (Lindenb.) Schiffn. Shoot (a): leaf lobule (b); leaf lobe (c); 
underleaf (d); female bract (e); cells of midleaf (f); female bracteole (g). Drawn from Meijer 387c, BO. 


Plants up to | cm long, 0.5-0.7 mm wide, blackish brown in dried condition. 
Branching Lejeunea-type. Stem diameter 40-110 um Leaves imbricate, 
widely spreading. Lobe oblong-orbicular, 250-500 um long, 200-400 um 
wide, margin entire, apex rounded; cells of lobe thick-walled, dark brown, 
isodiametric to hexagonal, trigones triangular, intermediate thickening 
scarce; marginal cells 6-10 x 5-10 um, mid-leaf cells 16-22 x 7-20 um, basal 
cells 20-28 x 16-20 um; oil bodies not seen. Lobule ovate, 120-230 um long, 
80-150 um wide, inflated, apex obliquely truncate, plane, connected to the 
leaf lobe across only one single cell. Underleaves distant to contiguous, 
obdeltoid to orbicular, 120-200 um long, 150-190 um wide, margin entire, 


Lejeuneaceae subfamily Prychanthoideae 69 


apex rounded to almost truncate, insertion line shallowly curved. Androecia 
not seen. Gynoecia on short or long branches, innovations lacking, bract 
oblong. 510-700 pm long, 270-330 um wide, margin entire, apex obtuse to 
orbicular, sinus up to 2/3 of lobe length, bract lobule about the same length 
as the bract lobe: bracteole obdeltoid to orbicular 330-700 um long, 350- 
500 pm wide, margin entire. sometimes recurved, apex broadly rounded. 
Perianth exserted, obovate, 300-900 um long, 250-500 um wide. inflated, with 
4 keels, margins toothed. Sporophytes and asexual reproduction not seen. 


Distribution: Java, Sumatra. Borneo, Sulawesi, Moluccas, West Irian, Papua New 
Guinea, Australia, Peninsular Malaysia, Philippines, Indochina, India, Bhutan. 
Nepal, China, Japan, tropical Africa, tropical America. 


Specimens examined: INDONESIA. West Java, without locality. collector 
unknown (BO!): Bogor Bot. Garden, van Borssum Waalkes 122, Meijer 
B995, B368, 55d4, Verdoorn 242 (BO!): Telaga Warna, Puncak Pass. 1500 m, 
Verdoorn 64d, 64b (BO!): G. Pangrango, “Tjisaroea™, ca 1200 m, Verdoorn 
44a (BO!); G. Guntur, “Kawah Kamoedjan”, 1500-1700 m, Verdoorn 50c 
(BO!). 


Notes: Diagnostic characters of Lopholejeunea nigricans are: 1) ovate leaf 
lobules, apex of lobule connected to the leaf lobe across only one single cell, 
2) entire margin of female bract and bracteole, 3) female bract lobule of 
almost the same length as the bract lobe. 4) perianth long exserted beyond 
the bracts, with 4 toothed keels (rarely entire). This species was found 
at ca 200-1700 m. It is separated from other species of the genus by the 
combination of characters mentioned above. 


9. Lopholejeunea ceylanica Steph., Sp. Hepat.5:86 (1912).—Type: Sri Lanka. 
Horton Plain, Giesenhagen s.n. (G, lectotype designated by Verdoorn, 
1934). 

— Lopholejeunea levieri Schiffn., Ann. Bryol. 6: 134 (1933).—Type: Indonesia. 
Sumatra, Mt. Singalang. “inter Riccardiam hymenophylloideum™, 1878, 
Beccari s.n. (holotype, FH) — cf. Zhu and Gradstein (2005). 

—Lopholejeunea schiffneri Verd., Ann. Bryol. 6: 134 (1933).—Type: Indonesia. 
Java, Cibodas, “ad cortices,” Jan 1895, Massart 1546 p.p. (holotype, FH) — cf. 
Zhu and Gradstein (2005). 


Dioicous. Plants up to 1.5 cm long, about 0.8 mm wide: blackish brown in the 
dried condition. Branching Lejeuwnea-type. Stem diameter 90-120 um Leaves 
imbricate, widely spreading. Lobe ovate-orbicular, 450-870 um long, 350-750 
um wide, margin entire, apex rounded to obtuse: cells of lobe thick-walled. 


70 Gard. Bull. Singapore 62 (1) 2010 


200 um 


4 2 q y, 
Soe ie 3 _ ee ie 
ae =< 


9 


- Figure 9. Lopholejeunea ceylanica Steph. Shoot (a); underleaf (b); leaf lobule (c); leaf lobe (d); 
female bract (e); female bracteole (f); cells of midleaf (g). Drawn from Haerida 1479, BO. 


brown, hexagonal to isodiametric, trigones triangular, intermediate thickening 
scarce; marginal cells 8-11 x 4-8 um, mid-leaf cells 14-30 x 15-23 um, basal 
cells 27-32 x 22-25 um; oil bodies not seen. Lobule oblong 250-450 um long, 
200-300 um wide, inflated, apex truncate with | small, unicellular tooth 
pointing towards the leaf apex, connected to the leaf lobe across 2-3 cells. 
Underleaves distant, orbicular, 150-300 um long, 190-350 um wide, margin 
entire, apex rounded, insertion line nearly straight. Androecia intercalary 
on branches, bracts in 5-9 pairs, similar to leaves but lobules larger, 400- 
430 um long, 250-350 um wide, bract lobule about 2/3 of the bract lobe, 
bracteoles similar to underleaves in size and shape. Gynoecia on short 
or long branches, innovations lacking, bract oblong to broadly obovate 
1000-1370 um long, 1050-1200 um wide, margin crenate, apex truncate to 
orbicular, bract lobule large, about the same length as the bract lobe, oblong, 
margin entire, apex acute; bracteole obovate 900-1200 um long, 1050-1100 
um wide, margin entire, crenulate towards the apex, apex rounded. Perianth 
immersed, obovate, 850 um long, 630 um wide, inflated, with 4 keels, margins 
toothed. Sporophytes and asexual reproduction not seen. 


Specimens examined: INDONESIA. West Java, Telaga Warna, Haerida 1479 
(BO!); Cibodas Bot. Garden, ca 1450 m, Meijer B3801, 3810, Neervoort 811, 
3353 (BO!);G. Pangrango, tea estate Mandalawangi, ca 1600 m, Meijer B387g 
(BO!); G. Gede, Kandang Badak 2400 m, subalpine forest (forest canopy 
ca 10 m high), common on tree trunks, Gradstein10241 (BIOT!, GOET!); 


Lejeuneaceae subfamily Ptychanthoideae 7h 


G. Cikurai, W slope, ca 1700 m, Verdoorn 59] (BO!); Cigombong (Pondok 
Gedelanden), ca 500 m, Verdoorn 43c (BO!): G. Patuha, Kornochalert 1405 
(BIOT!). 


Distribution: Java, Sumatra, Borneo, Peninsular Malaysia, Sri Lanka, 
Indochina. 


Notes: Diagnostic characters of Lopholejeunea ceylanica are: 1) lobule large, 
ca ¥2 x leaf length, with one small, unicellular tooth pointing towards the 
leaf apex, 2) apex of leaf- lobule attached to the lobe across 2-3 cells, 3) 
crenate margin of female bract, 4) female bract lobule large, acute. This 
species grows at ca 500-2400 m. It resembles the very common L. subfusca 
but differs by the more brownish color, the larger lobule (ca % x leaf length; 
in L. subfusca ca 1/3 x leaf length) and the large lobule of the female bract 
(very small in L. subfusca). 


10. Lopholejeunea horticola Schiffn., Ann. Bryol. 6: 133 (1933). —Type: 
Indonesia. Java, Bogor Bot. Garden, on trees (“In horto Buitenzorgensi ad 
arbores”), Dec 1894, Massart 941 (holotype, FH; isotypes, FH, W) — cf. Zhu 
and Gradstein (2005). 

— Lopholejeunea glomerata Herzog, Ann. Nat. Mus. Wien 53: 370 (1942). 
—Type: Indonesia. Sumatra, Padang, Schild 119 (holotype, W) — cf. Zhu and 
Gradstein (2005). 


( ie ; a 100 Hm) \ 


Figure 10. Lopholejeunea horticola Schiffn. Shoot (a); underleaf (b); leaf lobule (c); female bract 
(d), female bracteole (e); cells of midleaf (f); cells of margin of leaf (g); perianth (h). Drawn from 
Haerida 809, BO. 


Te Gard. Bull. Singapore 62 (1) 2010 


Dioicous. Plants up to 2 cm long, 0.6-0.9 mm wide, dark brown in dried 
condition. Branching Lejeunea-type. Stem diameter 90-190 um Leaves 
imbricate, obliquely spreading. Lobe orbicular, oblong to quadrate, 450-800 
um long, 270-600 um wide, margin entire, apex rounded to obtuse; cells of 
lobe thick-walled, dark brown, hexagonal, trigones triangular, intermediate 
thickening scarce; marginal cells 7-10 x 6-8 um, mid-leaf cells 17-27 x 11-23 
um, basal cells 28-36 x 18-30 um; oil bodies not seen. Lobule ovate, 190- 
360 um long, 120-200 um wide, inflated, apex obliquely truncate, plane, 
sometimes with 1 small tooth, connected to the leaf lobe across 2-3 cells. 
Underleaves distant to contiguous, broadly-orbicular, 200-350 um long, 330- 
610 um wide, margin entire, bases cuneate, apex rounded to almost truncate, 
insertion line nearly straight. Androecia intercalary on branches, bracts in 
3-11 pairs, similar to leaves but lobules larger, 190-230 um long, 140-180 
um wide, bract lobule 2/3 of the bract lobe, strongly inflated, bracteoles 
similar to underleaves in size and shape, 100-180 um long, 120-200 um wide. 
Gynoecia on short or long branches, innovations lacking, bract obovate or 
_ oblong + 1200 um long, 900 um wide, margin entire, apex obtuse to truncate, 
bract lobule large, about the same length as the bract lobe, oblong, margin 
entire, apex obliquely truncate; bracteole obovate, about 800 um long, 900 
um wide, margin entire, apex truncate. Perianth immersed, obovate, about 
800 um long, 570 um wide, inflated, with 4 keels, margins strongly toothed. 
Sporophytes and asexual reproduction not seen. 


Specimens examined: INDONESIA. West Java, Cibodas Bot. Garden, 
Neervoort 2098 (BO!):G. Gede, Cibeureum, above “Soekaboemi”, 1200-1400 
m, Verdoorn 47b, 47a, 46 c (BO!); G. Gede Pangrango Nat. Park, Bodogol, 
Haerida 809 (BO!); G. Malabar, SW slope, ca 1650 m, Verdoorn 62a (BO!): 
G. Guntur, “Kawah Kamoedjan”, ca 1500-1700 m, Verdoorn 50a (BO!);: 
G. Gegerbentang, E slopes, ca 1500-2000 m, Verdoorn 67h (BO!); without 
locality, Pondok, Kurz s.n. (BO!); G. Patuha, ca 2000-2400 m, Verdoorn 60a 
(BO!). 


Distribution: Java, Sumatra, Bali, Moluccas, Peninsular Malaysia, Thailand. 


Notes: Diagnostic characters of Lopholejeunea horticola are: 1) minute plant, 
less than | mm wide, 2) lobule apex connected to the leaf lobe across 2-3 
cells, 3) female bract lobule almost as large as the lobe, 4) margins of female 
bract entire, 5) immersed perianth. This species grows at ca 800-2400 m. It 
closely resembles the very common L. subfusca but differs by the much 
smaller plant size and the large female bract lobule (very small in L. swbfusca). 


[e) 


Lejeuneaceae subfamily Prychanthoideae 7 


11. Lopholejeunea recurvata Mizut., J. Hattori Bot. Lab. 46: 369 (1979). 
—Type: Indonesia. Java, Gn. Gede, Cibodas, 1420 m, on bark of trees, Jun 
1930, F Verdoorn s.n. (holotype, NICH; isotypes, FH, JE, L. S. U) — cf. Zhu 
and Gradstein (2005). 


20um 


Figure 11. Lopholejeunea recurvata Mizut. Shoot (a): leaf lobe (b); leaf lobule (c); cells of midleaf 
(d). Drawn from Verdoorn 20f. BO. 


Autoicous. Plants up to 1.5 cm long, 1-1.2 mm wide; dark brown in dried 
condition. Branching Lejeunea-type. Stem diameter 60-130 um Leaves 
imbricate, widely spreading. Lobe oblong-rectangular, 430-760 um long, 
390-530 um wide, margin entire, apex truncate: cells of lobe thick-walled, 
brownish-orange, hexagonal toisodiametric,trigones triangular,intermediate 
thickening scarce; marginal cells 6-11 x 6-9 um, mid-leaf cells 13-20 x 10-17 
um, basal cells 26-33 x 11-19 um; oil bodies not seen. Lobule ovate, 160-220 
um long, 130-180 um wide, inflated, apex obliquely truncate, plane, connected 
to the leaf lobe across 2-3 cells. Underleaves contiguous to imbricate, wider 
than long, 150-250 um long, 220-400 um wide, margin entire, apex recurved, 
rounded to almost truncate, insertion line curved. Androecia intercalary on 
branches, bracts in 4-8 pairs, similar to leaves but lobules larger, 200-250 
um long, 100-150 um wide, bract lobule 1/2-2/3 of the bract lobe, bracteoles 
similar to underleaves in size and shape. Gynoecia not seen. 


Specimen examined: INDONESIA. West Java, Cibodas Bot. Garden, 
Verdoorn 20f (BO!). 


Distribution: Java, Sumatra, Bali, Peninsular Malaysia, Papua New Guinea. 


74 Gard. Bull. Singapore 62 (1) 2010 


Notes: Diagnostic characters of Lopholejeunea recurvata are: 1) recurved 
margins of leaf lobes and underleaves, 2) lobule apex connected to the leaf 
lobe across 2-3 cells. This rare species grows at ca 500-1900 m. It differs from 
other Javanese species of the genus by the recurved leaf margins. 


12. Lopholejeunea subfusca (Nees) Schiffn., Bot. Jahrb. Syst. 23: 593 (1897); 
Jungermannia subfusca Nees, Enum. Pl. Crypt. Jav. 1: 36 (1830); Lejeunea 
subfusca (Nees) Nees & Mont., Ann. Sci. Nat., Bot., Sér. 2, 5: 61 (1836); 
Phragmicoma subfusca (Nees) Nees, Naturgesch. Eur. Leberm. 3:248 (1838); 
Symbiezidium subfuscum (Nees) Trevis., Mem. Reale Ist. Lombardo Sci., 
Cl. Sci. Mat., Ser. 3, 4: 403 (1877); Lopholejeunea sagraeana var. } subfusca 
(Nees) Schiffn., Consp. Hepat. Archip. Ind.: 294 (1898). —Type: Indonesia. 
Java, without locality, Blume s.n. (holotype, STR; isotypes, W, S) — cf. Zhu 
and Gradstein (2005). 


200 ym 


(es 


on / 
iS | bherita, 2008 : és rs 
i See 
* |200 Hm at 
b> 
: 


Figure 12. Lopholejeunea subfusca (Nees) Schiffn. Shoot (a); leaf lobule (b); female bract (c); 
perianth (d); cells of midleaf (e). Drawn from Haerida 850, BO. 


Autoicous. Plants up to 2 cm long, 1-1.3 mm wide, dark brown to black 
in dried condition. Branching Lejeuwnea-type. Stem diameter 90-120 um. 
Leaves imbricate, widely spreading. Lobe oblong-orbicular, 410-700 tm 
long, 420-620 um wide, margin entire, apex rounded; cells of lobe thick- 
walled, pale brown to orange, hexagonal to isodiametric, trigones triangular, 
intermediate thickening scarce; marginal cells 6-9 x 5-7 um, mid-leaf cells 17- 
24 x 15-20 um, basal cells 22-30 x 16-23 um; oil bodies not seen. Lobule ovate, 
small, 180-220 um long, 120-150 um wide, inflated, apex obliquely truncate, 
plane, connected to the leaf lobe across 2-4 cells. Underleaves distant to 


Lejeuneaceae subfamily Ptychanthoideae iS 


contiguous, broadly-orbicular, 220-400 um long, 340-610 um wide, margin 
entire, apex truncate, insertion line shallowly curved. Androecia intercalary 
on branches, bracts in 3-16 pairs, similar to leaves but lobules larger, 230-430 
um long, 170-320 um wide, bract lobule almost the same size of the bract 
lobe, bracteoles similar to underleaves in shape, 170-270 um long, 180-450 
um wide. Gynoecia on short or long branches, innovations lacking, bracts and 
bracteoles in 2 pairs, bract ovate to broadly oblong 700-850 um long, 700-860 
um wide, margin irregularly toothed, apex orbicular to nearly truncate, sinus 
up to 1/4 of lobe length, bract lobule very small; bracteole broadly orbicular 
170-280 um long, 180-450 um wide, margin entire, apex rounded. Perianth 
immersed, obovate, about 870 um long, 600 um wide, inflated, with 2 keels, 
margins toothed. Sporophytes and asexual reproduction not seen. 


Specimens examined: INDONESIA. West Java, Bogor Bot. Garden, Meijer 
92b, 26b, Verdoorn 124, 141, 249 (BO!): Cibodas Bot. Garden, Neervoort 952, 
Verdoorn 20¢, 20d, Schiffner 250 (BO!), ibid., Lee & Nova Indri 49 (BIOT!, 
UKMB)!), ibid., ca 1425 m, Meijer B4017, B3800 (BO!); G. Gede Pangrango 
Nat. Park, Bodogol, ca 800 m, Haerida 850, 844 (BO!); G. Pangrango, ravine 
above Kampong G. Mas, Meijer B616i (BO!); Geger Bentang, Neervoort 
1344 (BO!); G. Halimun Nat. Park, Citalahab, 1065 m, Radhiah Zakaria 
254c (BIOT!); G. Papandayan, Tji Paroegpoeg, ca 2300-2500 m, Verdoorn 
54b (BO!); G. Cikurai, W slope, ca 1700 m, Verdoorn 59i (BO!); G. Patuha, 
Kornochalert 1416, 1400 (BIOT!). 


Distribution: Java, Sumatra, Borneo, Bali, Sulawesi, Moluccas, Philippines, 
Papua New Guinea, Pacific, Peninsular Malaysia, Indochina, China, Japan, 
India, Sri Lanka, Nepal, tropical Africa, tropical America. 


Notes: Diagnostic characters of Lopholejeunea subfusca are: 1) lobule apex 
truncate, connected to the leaf lobe across 2-4 cells, 2) very small size of 
female bract lobule, 3) entire margin of female bracteole, 4) immersed 
perianth. This very common species is very grows at ca 200-2500 m. Like 
in several other Javanese species of Lopholejeunea, the lobule apex is 
connected to the leaf lobe surface across 2-4 cells; the flat leaves with rounded 
apex, the orbicular, distant underleaves, the rather small leaf lobule, and the 
characters of the gynoecium mentioned above, readily separate this species 
from its relatives. 


13. Lopholejeunea wiltensii Steph., Hedwigia 35: 112 (1896).—Type: Indonesia. 
Sumatra, Padang, A. Wiltens (holotype, G) — cf. Zhu and Gradstein (2005). 


76 Gard. Bull. Singapore 62 (1) 2010 


— Lopholejeunea serrifolia Steph., Sp. Hepat. 5: 84 (1912). —Type: Indonesia. 
Java, without locality, collector unknown (G, lectotype designated by 
Mizutani, 1979). 


i " 
200"um } Hara 2004 ih 
a : 


Figure 13. Lopholejeunea wiltensii Steph. Shoot (a); leaf lobule (b); cells of midleaf (c); female 
bracteole (d): female bract (e). Drawn from Verdoorn 30d, BO. 


Dioicous. Plants up to 4 cm long, 0.9-1.3 mm wide; pale brown in dried 
condition. Branching Lejeuwnea-type. Stem diameter 50-100 um Leaves 
imbricate, widely spreading. Lobe broadly ovate to broadly orbicular, 500- 
650 um long, 580-610 um wide, margin entire, apex rounded; cells of lobe 
thick-walled, pale yellow, quadrate to isodiametric, trigones triangular, 
intermediate thickening scarce; marginal cells 5-7 x 2-6 um, mid-leaf cells 
11-13 x 7-10 um, basal cells 20-25 x 13-23 um; oil bodies not seen. Lobule 
triangular to quadrate, small, 190-220 um long, 110-130 um wide, inflated 
at the basal part forming somewhat a longitudinally elliptical sac, apex 
obliquely truncate with 1 tooth consisting of 1-2 cells, connected to the leaf 
lobe across 2-3 cells. Underleaves distant to contiguous, reniform, 210-360 
um long, 310-610 um wide, margin entire, apex rounded to almost truncate, 
insertion line strongly curved. Androecia not seen. Gynoecia on short or 
long branches, innovations lacking, bract oblong-ovate to broadly ovate, 
about 470 um long, 460 um wide, margin irregularly toothed, apex obtuse to 
orbicular, sinus up to 1/3 of lobe length, bract lobule oblong about 1/3 of the 
bract lobe length, margin entire; bracteole broadly ovate about 350 um long, 
410 um wide, margin irregularly toothed, apex truncate. Perianth immersed, 
obovate, about 970 um long, 670 um wide, inflated, with 4 keels, margins 
toothed. Sporophytes and asexual reproduction not seen. 


Lejeuneaceae subfamily Ptychanthoideae al 


Distribution: Java, Sumatra, Peninsular Malaysia, Philippines, Sulawesi. 
Moluccas, West Irian, Papua New Guinea, New Caledonia. 


Specimens examined: West Java, Cibodas Bot. Garden, 1400 m, Ariyanti 458 
(BIOT!), ibid., along the road to Huis ten Bosch, Meijer B1078 (BO!): above 
Cibodas, Cibeureum, Schiffner 252, 253, Verdoorn 30d, 30c, 30a (BO!); G. 
Gede, Soekaboemi, Verdoorn 46a (BO!): G. Pangrango, *Tjisaroea”, ca 1200 
m, Verdoorn 44a (BO!):Tugu, tea estate G. Mas, Meijer & van der Wijk BO59b 
(BO!); above tea estate G. Mas, Jalan Mandalawangi, Meijer B335a (BO!): 
G. Guntur, “Kawah Kamoedjan”, ca 1500-1700 m, Verdoorn 50b (BO!). 


Notes: Diagnostic characters of Lopholejeunea wiltensii are: 1) pale color of 
the plant in the dried condition, 2) lobule small, in the basal part forming a 
somewhat longitudinally elliptical sac, 3) lobule apex connected to the leaf 
lobe across 2-3 cells, 4) reniform underleaves. This species grows at ca 1200- 
1900 m. Lopholejeunea wiltensii is closely related to L. zollingeri and was 
sometimes considered a synonym of the latter. It differs from L. zollingeri 
by the characters given in the key. 


14. Lopholejeunea zollingeri (Steph.) Schiffn.,Consp. Hepat. Arch. Ind.:296 
(1898); Lejeunea zollingeri Steph., Hedwigia 29: 14 (1890). —-Type: Indonesia. 
Java, without locality, collector unknown (W, lectotype designated by 
Mizutani, 1979). 

— Lopholejeunea latialata Mizut., J. Hattori Bot. Lab. 46: 365 (1979). —Type: 
Indonesia. Java: without locality, collector unknown, ex Herb. Sande Lacoste 
(holotype, L) — cf. Zhu and Gradstein (2005). 


7 


Ye 
hy es 


Figure 14. Lopholejeunea zollingeri (Steph.) Schiffn. Shoot (a): leaf lobule (b): leaf lobe (c): 
underleaf (d); cells of midleaf (e). Drawn from Meijer 387c, BO. 


78 Gard. Bull. Singapore 62 (1) 2010 


Autoicous. Plants up to 3 cm long, 0.7-1.2 mm wide, blackish brown in dried 
condition. Branching Lejeunea-type. Stem diameter 60-160 um. Leaves 
imbricate, widely spreading. Lobe orbicular to broadly ovate, 200-420 um 
long, 230-460 um wide, margin entire, apex rounded, usually recurved; cells 
of lobe thick-walled, brown, hexagonal to isodiametric, trigones triangular, 
intermediate thickening frequent; marginal cells 5-11 x 7-9 um, mid-leaf cells 
11-19 x 9-15 um, basal cells 25-30 x 19-22 um; oil bodies not seen. Lobule 
ovate, large, almost 1/3 of the leaf lobes length, 190-210 um long, 160-180 um 
wide, inflated at the middle part forming a globose sac, apex obliquely truncate 
with | small tooth or without tooth but forming a sharp angle, connected to 
the leaf lobe across 3-4 cells. Underleaves closely imbricate, reniform, 190-420 
um long, 370-710 um wide, margin entire, apex rounded to almost truncate, 
insertion line strongly arched. Androecia intercalary on branches, bracts in 4- 
9 pairs, similar to leaves but lobules larger, 200-220 um long, 160-190 um wide, 
bract lobule 1/3 of the bract lobe, bracteoles similar to underleaves in shape, 
110-170 um long, 180-300 um wide. Gynoecia not seen. 


Specimens examined: INDONESIA. West Java, G. Pangrango, tea estate 
Mandalawangi, ca 1600 m, Meijer B387c (BO!); G. Patuha, ca 1600 m, 
Kornchalert 1413, Gradstein 12169 (BIOT!). 


Distribution: Java, Sumatra, Borneo, Sulawesi, Peninsular Malaysia, 
Philippines, Papua New Guinea, Sri Lanka, China, Japan, Fiji. 


Notes: Diagnostic characters of Lopholejeunea zollingeri are: 1) broadly 
orbicular leaf lobe with recurved apex, 2) leaf lobule large, apex connected 
to the leaf lobe across 3-4 cells, 3) closely imbricate, large, reniform 
underleaves with strongly arched insertion line. This species grows at ca 
1350-1600 m. Lopholejeunea zollingeri has very large, reniform underleaves 
like in L. wiltensti, L. eulopha and L. applanata but differs from the latter 
two by the lobule apex which is attached to the lobe across 3-4 cells (across 
1 cell in L. eulopha and L. applanata). For differences with L. wiltensii see 
the characters given in the key. 


15. Mastigolejeunea auriculata (Wils.) Schiffn., in Engler and Prantl, Nat. 
Pflanzenfam. 1, 3: 129 (1893); Jungermannia auriculata Wils., in Drummond, 
Musci Amer. Exsicc. (Southern States) nr. 170 (1841); Ptychocoleus 
auriculatus (Wils.) Trevis., Mem. Reale Ist. Sci. Mat. Nat., Ser. 3, 4: 405 
(1877). -Type: USA. Louisiana, New Orleans, Drummond s.n. (holotype, 
BM; isotypes, MANCH, PC) —cf. Gradstein et al. (2002). 

—Phragmicoma humilis Gottsche, in Gottsche et al., Syn. Hepat. 299 (1845); 
Mastigolejeunea humilis (Gottsche) Schiffn., in Engler and Prantl, Nat. 


Lejeuneaceae subfamily Ptychanthoideae 79 


Pflanzenfam. 1, 3: 129 (1893). —Type: Indonesia. Java, without locality, 
Junghuhn s.n. (isotypes, B, W) — cf. Gradstein et al. (2002). 


Figure 15. Mastigolejeunea auriculata (Wils.) Schiffn. Shoot (a); underleaf (b):; leaf lobe (c); leaf 
lobule (d); cells of basal part of the leaf (e); cells of midleaf (f). Drawn from Meijer 92a, BO. 


Plants up to 2 cm long, 0.8-1 mm wide, dull green to dark brown in dried 
condition. Branching Lejeunea-type. Stem diameter 70-170 um Leaves 
imbricate, obliquely spreading, somewhat squarrose. Lobe ovate, 440-1030 
um long, 410-730 um wide, margin entire, apex rounded to obtuse; cells of 
lobe thick-walled, light yellow, rhomboidal to hexagonal, trigones cordate, 
intermediate thickening scarce; marginal cells 11-15 x 6-10 um, mid-leaf cells 
18-21 x 9-12 um, basal cells 25-34 x 16-19 um; oil bodies not seen. Lobule 
oblong to subrectangular, 190-440 um long, 130-200 um wide, apex truncate 
to obliquely truncate with 1 small tooth. Underleaves imbricate, obdeltoid 
to nearly triangular, 200-530 um long, 220-490 um wide, margin entire, apex 
truncate, base cuneate, insertion line curved. Androecia not seen. Gynoecia 
on short or long branches, with 1-2 lejeuneoid innovations, bract oblong to 
broadly oblong, 1000-1070 um long, 600-800 um wide, margin entire, apex 
subrounded to nearly truncate, sinus 1/3 to 2/3 of lobe length, bract lobule 
oblong-ovate to subrectangular, 500-670 um long, 340-360 um wide, margin 
somewhat undulated towards the apex; bracteole obdeltoid 600-770 um 
long, 850-960 um wide, margin entire, apex truncate. Perianth obovate, 1010 
um long, 440 um wide, with 3 keels, margins entire. Asexual reproduction 
not observed. 


Specimens examined: INDONESIA. West Java, Bogor Bot. Garden, van 
Borssum Waalkes 122a, Meijer 92a, 552d (BO!). Banten, Taman Nasional 
Ujung Kulon, Dewi Dw 922b (BO!). 


80 Gard. Bull. Singapore 62 (1) 2010 


Distribution: Java, Borneo, Sulawesi, Moluccas, West Irian, Papua New 
Guinea, Australia, Solomon Is., India, Indochina, China, Japan, tropical 
Africa, tropical America. 


Notes: Diagnostic characters of Mastigolejeunea auriculata are: 1) leaf lobe 
ovate in shape, margins entire, 2) leaf cells elongate, with cordate trigones 
3) lobule with truncate to obliquely truncate apex, with 1 small tooth, 4) 
underleaves obdeltoid, 5) perianth 3-keeled, with smooth keels and 1-2 
innovations. This species grows at ca 200-1700 m. It is very similar to M. 
virens but differs by the very short, blunt lobule tooth. 


16. Mastigolejeunea indica Steph., Sp. Hepat. 4: 776 (1912). —Type: India. 
Nicobar Is., Man s.n. (holotype, G) —cf. Gradstein et al. (2002). 
—Thysananthus integrifolius Steph., Sp. Hepat. 4: 788 (1912). Mastigolejeunea 
integrifolia (Steph.) Verdoorn, Blumea 1: 231, 239 (1934). —Type: Australia. 
Torres Str., Possession I., Micholitz s.n. (holotype, G; isotype, FH) -cf. 
Gradstein et al. (2002). 


™ - 
ee 


Figure 16. Mastigolejeunea indica Steph. Shoot (a); leaf lobe (b); underleaf (c); leaf lobule (d); cells 
of margin of leaf (e); cells of midleaf (f). Drawn from Meijer 75a, BO. 


Dioicous. Plants up to 2.5 cm long, 0.6-1 mm wide, 1-1.5 mm wide in wet 
condition, greenish-brown in dried condition, becoming more green towards 
the apex of the plant. Branching Lejewnea-type. Stem diameter 100-250 um, 
epidermis cells very unequal in size, dorsal one much large than ventral ones, 
hyalodermis lacking. Leaves imbricate, obliquely spreading, convolute when 


Lejeuneaceae subfamily Ptychanthoideae 8] 


dry. Lobe ovate-oblong, 520-1150 um long, 300-890 um wide, margin entire, 
apex acute to obtuse; cells of lobe thin-walled, light yellow, rhomboidal, 
trigones cordate, intermediate thickening scarce; marginal cells 6-11 x 3-5 
um, mid-leaf cells 7-18 x 6-14 um, basal cells 21-35 x 17-20 um; oil bodies not 
seen. Lobules ovate to triangular, 180-390 um long, 160-280 um wide, slightly 
inflated along the keel, apex obliquely truncate with 3-4 triangular teeth, each 
tooth consisting of 3-4 cells, 2-3 cells long, first and second tooth sometimes 
blunt, only 1-2 cells long, sometimes with or without fourth, 1-2 cells long 
tooth. Underleaves imbricate, obdeltoid, 230-520 um long, 250-540 um wide, 
margin entire, apex truncate, usually recurved and seemingly emarginate, 
base auriculate, insertion line curved. Generative structures not seen. 


Specimens examined: Inodnesia. West Java, Bogor Bot. Garden, ca 250m, 
Meijer 75a (BO!). 


Distribution: Java, China, India (Nicobar), Philippines, Papua New Guinea, 
Australia. 


Notes: Diagnostic characters of Mastigolejeunea indica are: 1) plant like 
M. auriculata but lobule with 3-4 large teeth, 2) stem cross section with 
very unequal epidermis cells, dorsal ones much large than ventral ones 
(hyalodermis lacking). Mastigolejeunea indica is new to Java. The species 
has only been collected in the Botanical Garden of Bogor, at ca 250 m. It 
is readily separated from all other species of the genus Mastigolejeunea by 
the lobule with 3-4 large triangular teeth (usually only one tooth in other 
species of the genus). 


17. Mastigolejeunea virens (Angstr.) Steph., in Sp. vee A/G. VOID: 
Thysananthus virens Angstr., Ofv. K.Vetensk Akad. Férh. 5: 131. 1873. - 
Type: Society Is., Moorea, Andersson s.n. (holotype, S; ieee FH, G) -cf. 
Gradstein et al. (2002). 

—Mastigolejeunea humilis sensu Verdoorn 1933 p.p. 


Autoicous. Plants up to 3 cm long, 0.9-2 mm wide, brown to dark brown 
in dried condition. Branching Lejeunea-type. Stem diameter 60-190 um 
Leaves closely imbricate, widely spreading. Lobe ovate to oblong, 340-920 
um long, 200-640 um wide, margin entire, apex rounded to obtuse; cells of 
lobe thick-walled, light-yellow, rhomboidal, trigones cordate, intermediate 
thickening scarce; marginal cells 6-11 x 5-6 um, mid-leaf cells 11-16 x 5- 
7 um, basal cells 18-26 x 8-10 um; oil bodies not seen. Lobule ovate, 140- 
280 um long, 60-170 um wide, inflated, apex obliquely truncate with 1 long 
tooth consisting of 3-5 cells. Underleaves imbricate, quadrangular to nearly 


82 Gard. Bull. Singapore 62 (1) 2010 


triangular, 190-560 um long, 250-600 um wide, margin entire, apex truncate, 
sometimes recurved forming a hearth shaped. Androecia intercalary on 
branches, bracts in 5 pairs, similar to leaves but lobules larger, bract lobule 
2/3 of the bract lobe, 100-110 um long, 50-60 um wide, bracteoles similar to 
underleaves in size and shape, 90-120 um long, 100-120 um wide. Gynoecium 
with 1-2 subfloral innovations; innovation type lejeuneoid; bract lobe ovate- 
oblong, ventral margin incurved, margin entire, apex rounded, 1160 um 
long, 460 um wide; bract lobule about % of bract lobe length, oblong, margin 
crenulate; bracteole oblong, margin entire, apex rounded. Perianth 820 um 
long, oblong, inflated, with 3 keels, margins entire. Sporophytes and asexual 
reproduction not seen. 


10 Hm 


Figure 17. Mastigolejeunea virens (Angstr.) Steph. Shoot (a); leaf lobe (b); underleaf (c); cells of 
midleaf (d); cells of basal part of the leaf (e). Drawn from Meijer 62a, BO. 


Specimens examined: INDONESIA. West Java, Bogor, without locality, 
Verdoorn 149 (BO!); Bogor Bot. Garden, Schiffner 257, W. Meijer B24, 
62a, B55f, B998b, Verdoorn 12z, 12d, 143, 255 (BO!); Cibodas Bot. Garden, 
Neervoort 154, 918, Meijer B3720, B3815 (BO!), ibid., Lee & Nova Indri 62 
(BIOT!, UKMB!), ibid., Ariyanti 472 (BIOT!); Telaga Bodas, Verdoorn 58a, 
5Sb (BO!); G. Megamendung, above “Toegoe”, Schiffner 256 (BO!). 


Distribution: Java, Borneo, Moluccas, Philippines, Peninsular Malaysia, 
Thailand, Sri Lanka, Papua New Guinea, Australia, Pacific Islands. 


Notes: The occurrence of Mastigolejeunea virens in Java has not been 
reported before, therefore this species is new for Java. Verdoorn (1933) 


Lejeuneaceae subfamily Ptychanthoideae 83 


treated the species as a synonym of M. humilis (= M. auriculata), but it is 
readily separated from the latter by the longer lobule tooth (see key). The 
diagnostic characters of Mastigolejeunea virens are: 1) leaf lobe rounded to 
obtuse at apex, 2) lobule with | long tooth consisting of 3-5 cells, 3) perianth 
with 3 keels. This species grows at ca 200-1500 m. 


18. Ptychanthus striatus (Lehm. & Lindenb.) Nees, Naturgesch. Eur. 
Leberm. 3: 212 (1838); Jungermannia striata Lehm. & Lindenb., Nov. Stirp. 
Pug. 4: 16 (1832); Bryopteris striata (Lehm. & Lindenb.) Mitt., in Seemann, 
Fl. Vit. 411 (1873). -Type: Nepal. Wallich s.n. & s.d. (holotype, S; isotypes, G, 
W) -cf. Gradstein er al. (2002). 

—Jungermannia retusa Reinw. et al. var. f Nees, Enum. PI. Crypt. Javae 1: 39 
(1830); Ptychanthus retusus (Reinw. et al.) Nees var. B Nees, in Gottsche et al., 
Syn. Hepat. 292. 1845: Ptycholejeunea retusa (Reinw. et al.) Steph., Hedwigia 
28: 258 (1889); Ptychanthus retusus (Reinw. et al.) Steph., Sp. Hepat. 4: 743 
(1912): Ptychanthus striatus var. retusus (Reinw. et al.) Verd., Ann. Bryol. 
Suppl. 4: 122 (1934). —Type: Indonesia. Java, without locality, Blume s.n. 
(holotype, STR; isotype, W) —cf. Gradstein et al. (2002). 


Figure 18. Prychanthus striatus (Lehm. & Lindenb.) Nees. Shoot (a); underleaf (b): leaf lobe (c): 
cells of midleaf (d). Drawn from Meijer 840, BO. 


Autoicous. Plants robust, up to 10 cm long, 1-1.5 mm wide, standing up away 
from the substrate or pendent, pinnate, greenish-brown in dried condition. 
Branching Frullania-type. Stem diameter 130-300 um Leaves imbricate, 
flat, obliquely spreading. Lobe broadly-ovate, 650-2170 um long, 300-1260 
um wide, margin entire or slightly toothed towards the apex, apex acute, 
base auriculate at the proximal side; cells of lobe thick-walled, light-yellow, 


84 Gard. Bull. Singapore 62 (1) 2010 


rhomboidal to hexagonal, trigones cordate, intermediate thickening frequent; 
marginal cells 7-16 x 4-11 um, mid-leaf cells 20-27 x 8-14 um, basal cells 26-38 
x 11-20 um; oil bodies not seen. Lobule ovate, small, 240-370 um long, 110- 
170 um wide, inflated, apex with 1-2 teeth, teeth consisting of 1 small cell. 
Underleaves distant to contiguous, broadly-ovate, sometimes subrectangular 
in outline, 480-760 um long, 470-1000 um wide, base auriculate, margin entire, 
apex irregularly toothed. Androecia intercalary on branches, bracts in 5-10 
pairs, bract lobe ovate, 560-1150 um long, 300-520 um wide, margin entire, 
apex acute, bract lobule 1/2 of the bract lobe, inflated, 390-500 um long, 200- 
240 um wide, bracteoles similar to underleaves in size and shape. Gynoecia 
on short or long branches, innovation type lejeuneoid, bract lobe ovate with 
small bract lobules, 1220-1320 um long, 550-630 um wide, margin toothed 
towards the apex, apex acute, bract lobules oblong, 460-710 um long, 80- 
150 um wide; bracteole broadly ovate, 1110-1150 um long, 1220-1330 um 
wide, margin toothed, apex rounded. Perianth immersed, elliptical, 1770 um 
long, 730 um wide, inflated, with + 10 keels, margins entire. Sporophytes and 
- asexual reproduction not seen. 


Specimens examined: INDONESIA. West Java, G. Salak, ca 1000 m, 
Zollinger 3560, Schiffner 259 (BO!); G. Gede, above Cibodas, 1600-1900 m. 
Neervoort 107, 129, 227, 2775, 2401, 282, 2271, Verdoorn 30g, 30f, 21a, 30n, 
Iwamasa s.n., Meijer B4137, B4076, B160 (BO!), ibid., above Tanjung Mas 
waterfall (Pancuran Mas), Alston 12807 (BO!), ibid., Cibeureum, Schiffner 
266, Verdoorn 30h, 301, 30}, 30n (BO!), ibid., trail Cibeureum falls to Kandang 
Badak 1700-2200 m, montane forest, epiphytes on trunk bases, common, 
Gradstein 10215, 10217 (BIOT!, GOET!); G. Pangrango, “Tjisaroea”, ca 1200 
m, Verdoorn 44a, 44b, 44c (BO!); Tugu, slope of Gede-Pangrango, above G. 
Mas, Meijer B3394a, BS40, B421 (BO!); G. Geger Bentang, E slopes, 1500- 
2000 m, Neervoort 2890, Verdoorn 67a, 67b, 267, Meijer B5584 (BO!); G. 
Megamendung, Schiffner 265 (BO!); G. Malabar, SW slopes of Punciak 
Besar 1650-2300 m, Verdoorn 61a, 61b, 61c, 62b, 62d (BO!); G. Patuha, 2000- 
2400 m, Verdoorn 60b (BO!), Kornochalert 1414 (BIOT!). 


Distribution: Java,Sumatra, Peninsular Malaysia, Borneo,Sulawesi, Moluccas, 
Philippines, West Irian, Papua New Guinea, Australia, New Zealand, Pacific 
Islands, India, Sri Lanka, Taiwan, Indochina, China, Japan, tropical Africa. 


Notes: Diagnostic characters of Ptychanthus striatus are: 1) Frullania-type 
branching, 2) lobe with acute apex and entire or toothed margins, 3) leaf 
cells elongate, with cordate trigones, 4) perianth with 10 smooth keels, and 
with innovation, 5) plant robust, pinnate, usually pendent. This species is 
common in the mountains where it occurs at ca 1000-2400 m. By its large 


(oe) 
Nn 


Lejeuneaceae subfamily Ptychanthoideae 


size, its pinnate, Frullania-type branching and acute leaf lobes the speciers 
is unmistakable and cannot be confused with any other Javanese member 
of Lejeuneaceae. 


19. Schiffneriolejeunea pulopenangensis (Gottsche) Gradst., J. Hattori Bot. 
Lab. 38: 335 (1974); Phragmicoma pulopenangensis Gottsche, Syn. Hepat. 
299 (1845); Ptychocoleus pulopenangensis (Gottsche) Trevis., Mem. Reale Ist. 
Lombardo Sci., Cl. Sci. Mat., Ser. 3, 4: 405 (1877).—Type: Peninsular Malaysia. 
Pulo Penang, Delessert s.n. (holotype, PC-Mont; isotypes, BM, S, W). 


Plants up to6cm long, 1.5-2 mm wide, brown to dark brown in dried condition. 
Vegetative branching Frullania-type, or Lejeunea-type. Stem diameter 150- 
260 um. Leaves imbricate, obliquely spreading, convolute when dry. Lobe 
broadly ovate to somewhat triangular, 700-1060 um long, 500-850 um wide, 
margin entire, apex obtuse to nearly acute; cells of lobe thick-walled, hyaline, 
rhomboidal to hexagonal, trigones cordate, intermediate thickening scarce; 
marginal cells 9-24 x 7-16 um, mid-leaf cells 17-26 x 8-19 um, basal cells 25- 
45 x 13-23 um; oil bodies not seen. Lobule ovate, 300-420 um long, 150-280 


100pHm 


Figure 19. Schiffneriolejeunea pulopenangensis (Gottsche) Gradst. Shoot (a); leaf lobe (b); apex 
of leaf lobule (c); underleaf (d); cells of midleaf (e). Drawn from Meijer & Alston 5329, BO. 


um wide, apex with 2 conspicuous teeth that point outwards towards the 
leaf apex, first tooth consisting of 3-4 cells, second tooth consisting of 2- 
3 cells, lobule free margin plane, with 2 clearly visible teeth. Underleaves 
imbricate, obdeltoid-orbicular sometimes with recurved apex seemingly 
emarginate, 360-420 um long, 480-610 um wide, margin entire, apex truncate, 


86 Gard. Bull. Singapore 62 (1) 2010 


insertion line deeply curved. Androecia on lateral branches, inflated, in 3-11 
pairs, about 1/3-1/2 of the vegetative lobe, hypostatic, bracteoles oblong to 
obdeltoid, apex truncate, smaller than the underleaves. Gynoecia not seen. 


Specimen examined: INDONESIA. West Java, Bogor Bot. Garden, Meijer 
& Alston 5329 (BO!). 


Distribution: Indomalesia, Australia. 


Notes: Diagnostic characters of Schiffneriolejeunea pulopenangensis are: 
1) branching Lejeunea and Frullania type, 2) leaf lobule ovate, with plane 
free margin and with 2 long teeth that point outwards to the leaf apex, 3) 
leaves not squarrose when moist, convolute when dry, 4) trigones cordate. 
This species is apparently rare in Java and has only been collected in the 
Botanical Garden of Bogor at ca 250 m. The species is closely related to S. 
tumida but differs by the lobule without saccate base and by the flat leaves 
- (squarrose in S. tumida). 


20. Schiffneriolejeunea tumida var. haskarliana (Gottsche) Gradst. & 
Terken, Occas. Pap. Farlow Herb. 16: 77 (1981); Phragmicoma hasskarliana 
Gottsche,in Gottsche eral.,Syn. Hepat.299 (1845): Acrolejeunea hasskarliana 
(Gottsche) Schiffn.,in Engler and Prantl, Nat. Pflanzenfam. 1, 3: 129 (1893); 
Ptychocoleus hasskarliana (Gottsche) Steph., Sp. Hepat. 5: 44 (1912). —Type: 
Indonesia. Java, without locality, Hasskarl 20 (holotype, B; isotypes, G, S, W) 
— cf. Gradstein and Terken (1981). 

—Mastigolejeunea badia Gottsche ex Steph., Sp. Hepat. 4: 779 (1912). —Type: 
Solomon Is., Vanikoro, Lesson s.n. (isotypes, BM, FH) —cf. Gradstein and 
Terken (1981). 


Plants up to 6 cm long, 2-3 mm wide; brown to dark brown in the dried condition. 
Vegetative branching Frullania-type, or Lejeunea-type. Stem diameter 160-270 
um. Leaves imbricate, obliquely spreading, somewhat squarrose, convolute 
when dry. Lobe broadly ovate to somewhat triangular, 900-1150 um long, 700- 
1200 um wide, margin entire, apex obtuse to nearly acute; cells of lobe thick- 
walled, hyaline, rhomboidal to hexagonal, trigones cordate, intermediate 
thickening scarce; marginal cells 13-24 x 9-22 um, mid-leaf cells 19-29 x 18-27 
um, basal cells 25-40 = 18-32 um; oil bodies not seen. Lobule ovate-rectangular, 
300-620 tm long, 130-270 um wide, apex with 2 teeth, each tooth consisting 
of 3-4 cells, lobule free margin strongly involute, forming a sac at the base of 
the lobule. Underleaves imbricate, obdeltoid sometimes with recurved apex 
seemingly emarginate, 420-840 um long, 490-720 um wide, margin entire, apex 
truncate, insertion line deeply curved. Generative structures not seen. 


Lejeuneaceae subfamily Ptychanthoideae 87 


I.Meerite.2009 | Fa 
Imm 


Figure 20. Schiffneriolejeunea tumida var. haskarliana (Gottsche) Gradst. & Terken. Shoot (a): leaf 
lobe (b); apex of leaf lobule (c); cells of midleaf (d). Drawn from Haerida 724, BO. 


Specimens examined: INDONESIA. West Java, Cibodas Bot. Garden, 
Neervoort 2120 (BO!), ibid., on bark of Altingia excelsa,ca 1400 m, Gradstein 
10202 (BIOT!, GOET!); above Cibodas, trail to Cibeureum waterfall, on 
tree trunk, Lee & Nova Indri 58 (BIOT!, UKMB!); G. Halimun Nat. Park, 
Haerida 724 (BO!). 


Distribution: Java, Papua New Guinea, Solomon Is., Australia. 


Notes: Diagnostic characters of Schiffneriolejeunea tumida var. haskarliana 
are: 1) branching Lejeunea- and Frullania-type, 2) leaf lobule with 2 
conspicuous, erect teeth and with a sac at the base by the strongly involute 
free margin, 3) leaves somewhat squarrose when moist, convolute when dry, 
4) cordate trigones. This species grows at ca 1000-1500 m. It is closely related 
to S. pulopenangensis but occurs at higher elevations; for morphological 
differences see under the latter species and the key. 


21. Spruceanthus polymorphus (Sande Lac.) Verd., Ann. Bryol. Suppl. 4: 155 
(1934): Phragmicoma polymorpha Sande Lac., Ned. Kruidk. Arch. 34: 420 
(1854); Phragmolejeunea polymorpha (Sande Lac.) Schiffn., Forschungsr. 
Gazelle 4: 25. (1890); Thysananthus polymorphus (Sande Lac.) Schiffn., 
Consp. Hepat. Arch. Ind. 305 (1898): Archilejeunea polymorpha (Sande Lac.) 
B. Thiers & Gradst., Mem. N.Y. Bot. Garden 52: 10 (1989). —Type: Indonesia. 
Java, Junghuhn s.n. (holotype, L; isotype, NY) —cf. Gradstein et al. (2002). 


88 Gard. Bull. Singapore 62 (1) 2010 


Autoicous. Plants up to 4 cm long, 1.5-2.5 mm wide, pale green to greenish 
brown in dried condition. Branching Lejeunea-type. Stem diameter 100-250 
um: ventral merophyte 8-12 cells wide. Leaves imbricate, widely spreading. 
Lobe ovate-oblong, 730-1300 um long, 430-640 um wide, margins entire or 
toothed, plane or undulate, apex rounded to obtuse; cells of lobe isodiametric, 
with large, nodulose intermediate thickenings, hyaline, trigones not cordate: 
marginal cells 7-12 x 6-13 um, mid-leaf cells 20-26 x 19-24 um, basal cells 
26-52 x 13-18 um: oil bodies not seen. Lobule ovate, 110-230 um long, 80-130 
um wide, apex obliquely truncate with 1-2 small teeth. Underleaves closely 
imbricate, obdeltoid to orbicular, 230-400 um long, 400-500 um wide, margin 
entire at lateral part, serrate towards the apex, apex rounded to truncate, 
insertion line shallowly curved. Androecia intercalary on branches, bracts 
up to 14 pairs, similar to leaves but lobules larger, 580-650 um long, 250-300 
um wide, hypostatic, bracteoles similar to underleaves in size and shape. 
Gynoecia with 1-2 lejeuneoid innovations, bract ovate, about 900 um long, 
600 um wide, margin entire, apex rounded, bract lobule ovate about 2/3 of 
- the lobe length, toothed; bracteole orbicular to oblong, 530-600 um long, 
580-900 um wide, margin toothed. Perianth oblong, about 1300 um long, 
430 um wide, with 7 smooth keels, margins entire. Sporophytes and asexual 
reproduction not seen. 


20 um 


Figure 21. Spruceanthus polymorphus (Sande Lac.) Verd. Shoot (a); underleaf (b): cells of basal 
cells of the lobe (c); leaf lobe (d); cells of midleaf (e). Drawn from Borssum Waalkes 450, BO. 


Specimens examined: INDONESIA. West Java, Cibodas Bot. Garden, Meijer 
B3820, B178 (BO!), ibid., on bark of Araucaria, ca 1400 m, very common, 
Gradstein 10207 (BIOT! GOET!), ibid., near the guest house, on base of 


Lejeuneaceae subfamily Ptychanthoideae 89 


tree, Gradstein 10206 (BIOT! GOET!); Gn. Pangrango, Meijer B605 (BO!): 
Gn. Gede, 1500-1900 m, Verdoorn 46e, 46d, 47c, Meijer B589a2 (BO!), ibid., 
N slope above “Artja”, ca 1100 m, Schiffner 287, 285 (BO!): Rawa Denok, ca 
1900 m, Neervoort 2380 (BO!); Gn. Halimun, ca 930 m, collector unknown 
(BIOT!): Gn. Cikurai, ca 1700 m, Verdoorn 59a, 59b,59c, 59d, 59e, 59f (BO!): 
Geger Bentang, Neervoort 1366, 1089 (BO!); Gn. Malabar, Verdoorn 62c 
(BO!). Banten, Pulau Panaitan, Gn. Putri, ca 75 m, van Borssum Waalkes 
450 (BO!); Taman Nasional Ujung Kulon, Dewi Dw928, 922c (BO!). 
Distribution: Java, Sumatra, Borneo, Sulawesi, Philippines, New Guinea, 
Australia, Pacific, Japan, Taiwan, India. 


Notes: Diagnostic characters of Spruceanthus polymorphus are: 1) rather 
robust plant, ventral merophyte 8-12 cells wide, 2) underleaves serrate 
towards the apex, 3) leaf lobes rounded to obtuse at apex, with entire or 
toothed, plane or undulate margin, 4) leaf cells isodiametrical,5) female bract 
lobe entire, female bracteole toothed 6) perianth with up to 7 smooth keels, 
1-2 innovations present. This species grows at ca 75-1900 m. Spruceanthus 
polymorphus may be confused with S. semirepandus and Archilejeunea 
planiuscula. S. semirepandus differs by the acute leaf apex (rounded to 
obtuse in S. polymorphus) and A. planiuscula by the thinner stems and the 
segmented oil bodies (although we did not observe the oil bodies, they are 
supposedly homogeneous in S. polymorphus). 


22. Spruceanthus semirepandus (Nees) Verd., Ann. Bryol. Suppl. 4: 153 
(1934): Jungermannia semirepanda Nees, Enum. PI. Crypt. Javae 1:39 (1830); 
Ptychanthus semirepandus (Nees) Nees, Naturg. Eur. Leberm. 3: 212 (1838); 
Phragmicoma semirepanda (Nees) Gottsche, in Gottsche ef al., Syn. Hepat. 
302 (1845); Lejeunea semirepanda (Nees) Mitt., J. Proc. Linn. Soc. Bot. 5: 111 
(1861). —Type: Indonesia. Java. Blume s.n. (holotype, STR?) — cf. Verdoorn 
(1934). 


Autoicous. Plants robust, up to 8 cm long, 1.5-3 mm wide, pale brown to dark 
brownin dried condition. Branching Lejeunea-type.Stem diameter 120-300 um; 
ventral merophyte 8-12 cells wide. Leaves imbricate, widely spreading. Lobe 
ovate, 1000-2300 um long, 550-1590 um wide, margin entire, crenulate towards 
the sharply acute apex: cells of lobe thick-walled, light yellow, hexagonal 
and almost isodiametric, trigones not cordate, intermediate thickening 
frequent; marginal cells 5-15 x 4-10 um, mid-leaf cells 20-31 x 13-20 um, 
basal cells 30-41 x 21-30 um; oil bodies not seen. Lobule ovate, 240-450 
um long, 120-220 um wide, inflated, apex truncate with 1-2 small teeth. 
Underleaves imbricate, obdeltoid to rectangular, 500-1200 um long, 390-850 
um wide, margin entire, apex truncate, base auriculate. Androecia not seen. 


90 Gard. Bull. Singapore 62 (1) 2010 


Gynoecia with 1-2 lejeuneoid innovations, bracts ovate about 3050 um long, 
1530 um wide, margin toothed at the upper part, apex acute, bract lobule 
oblong, margin entire about 1/3 of the lobe lenght; bracteole broadly ovate 
to suborbicular with emarginate apex, toothed, about 2010 um long, 1500 
wide. Perianth, oblong-ovate, about 2700 um long, 1600 um wide, with 7- 
9 smooth keels, margins entire. Sporophytes and asexual reproduction not 
seen. 


Figure 22. Spruceanthus semirepandus (Nees) Verd. Shoot (a); underleaf (b); leaf lobe (c); leaf lobule 
(d); female bract (e); female bracteole (f); cells of midleaf (g). Drawn from Verdoorn 20c, BO. 


Specimens examined: INDONESIA. West Java, Cibodas Bot. Garden, 
Verdoorn 20c, 20b (BO!); Gn. Gede, Jwamasa s.n. (BO!); Gn. Malabar, 1800- 
2300 m, Verdoorn 61d (BO!); Gn. Cikurai, ca 1700 m, Verdoorn 59k (BO!); 
Gn. Patuha, ca 1500 m, Kornochalert 1417 (BIOT!). 


Distribution: Java, Borneo, Moluccas, Philippines, China, Taiwan, India, Sri 
Lanka, Japan. 


Notes: Diagnostic characters of Spruceanthus semirepandus are: 1) robust 
plant, ventral merophyte to 12 cells wide, 2) underleaves with entire 
margin, 3) leaf lobe acute at apex, coarsely dentate at margin, 4) leaf cells 
isodiametric 4) female bract lobule entire, 5) perianth with 7-9 smooth 
keels, innovation present. This species grows at 1200-2300 m. It is closely 
related to S. polymorphus but the leaf apex in the latter species is broader, 
rounded to obtuse. By its acute leaves S. semirepandus may be confused with 


Lejeuneaceae subfamily Ptychanthoideae 9] 


Ptychanthus striatus but the very different branching types (Lejeunea-type 
in Spruceanthus, Frullania-type in Ptychanthus) readily separates the two. 


23. Thysananthus convolutus Lindenb., in Gottsche et al., Syn. Hepat. 288 
(1845).—Type: Indonesia. Java, unknown locality, collector unknown.., ex hb. 
Lindenberg (isosyntype: G) — cf. Gradstein er al. (2002). 


Plants up to 5 cm long, 0.8-1.2 mm wide, pale brown to dark brown in 
dried condition. Branching Lejeunea-type. Stem diameter 100-280 um 
Leaves closely imbricate, attached to the stem at an angle approx. 45-65°. 
Lobe ovate with recurved apex, asymmetric, upper part of leaf distinctly 
ventrad, 700-1150 um long, 600-810 um wide, margin toothed or entire, apex 
rounded; cells of lobe thick-walled, sometimes nodulose, trigones cordate, 
intermediate thickening frequent; marginal cells 7-20 x 6-11 um, mid-leaf 
cells 22-40 x 11-13 um, basal cells 36-52 x 11-15 um: oil bodies not seen. 
Lobule ovate, 250-320 um long, 110-130 um wide, inflated, apex truncate 
with 1-2 teeth, first tooth consisting of 3-4 cells, second tooth small consisting 
of 1 cell, sometimes without first tooth. Underleaves imbricate, suboblong, 
500-720 um long, 290-430 um wide, margins usually toothed, apex truncate, 
recurved, base not auriculate, insertion line shallowly curved. Generative 
structures not seen. 


400 pm 
¥ Pe bin 
| Herts 200815 53 3 OL 
U to) 20”um 


Figure 23. Thysananthus convolutus Lindenb. Shoot (a): underleaves (b): leaf lobe (c): cell of 
midleaf (d). Drawn from Jwamasa 5435, BO. 


92 Gard. Bull. Singapore 62 (1) 2010 


Specimens examined: INDONESIA. West Java, without locality, Teysmann 
s.n. (BO!); Kampung Dawuan, 1390 m, Neervoort 898 (BO!); Cibodas Bot. 
Garden, Pasir Sintek, ca 1410 m, Neervoort 3368 (BO!), ibid., on bark of 
trees at garden entrance, abundant, Gradstein 10205, on bark of Araucaria, 
Gradstein 10203 (BIOT!, GOET!); above Cibodas, trail to Cibeureum 
waterfall, Lee & Nova Indri 45 (BIOT!, UKMB!); Cibeureum, ca 1600-1900 
m, Verdoorn 30k, 30q, 20L, 21b, Iwamasa 5435, Neervoort 195, 308, Dadi & 
Noerta 134, Meijer B3578, B3705, B3789, B3578, V. Schiffner 283, 282a (BO!), 
N.S. Ariyanti 472 (BIOT!); Gn. Gede, boven Soekaboem1, ca 1500-1900 m, 
Verdoorn 46c, 46b, 46a (BO!); Gede-Pangrango, Tugu, above G. Mas, along 
JI. Mandalawangi, Meijer B3374, B3325, B387f, B607 (BO!); Geger Bentang, 
1400-1540 m, Neervoort 1178, 1059, 1024, 1806, 1019, 988, 2987, 3005 (BO!); 
Rawa Panjang, Neervoort 1166 (BO!); Cihoerang, ca 1380-1460 m, Neervoort 
138, 2241, 2242, Meijer B3681, B3741 (BO!); Telaga Warna, Puncak Pass, 
“Kratermuurtje”, Verdoorn 64f, 641, 64h, 64e (BO!); Gn. Halimun-Salak Nat. 
Park, Gn. Kendeng, ca 1250 m, Radhiah Zakaria 221c (BIOT!); Gn. Malabar, 
- SW slopes of Puncak Besar, 1800-2300 m, Verdoorn 61h (BO!). 


Distribution: Java, Sumatra, Peninsular Malaysia, Borneo, Sulawesi, 
Moluccas, Papua New Guinea, Philippines, Solomon Is. 


Notes: Diagnostic characters of Thysananthus convolutus are: 1) asymmetric 
leaf lobe with upper part ventrad and recurved, margin toothed or entire 2) 
underleaves suboblong, margins usually toothed or crenulate, 3) lobule with 
2 teeth. This species grows at ca 1300-2300 m. Forms with toothed leaves 
may be confused with 7: spathulistipus but the leaves in the latter species are 
symmetrical while those of 7? convolutus are asymmetric. Forms with entire 
leaves may be confused with Mastigolejeunea but the dorsal epidermis cells 
in Mastigolejeunea are larger than the inner stem cells, while in Thysananthus 
the epidermis cells are not larger than the inner cells. 


24. Thysananthus minor Verd.,in Rec. Trav. Bot. Neerl. 30: 231 (1933).—Type: 
Indonesia. Sumatra, Brastagi, Petani Falls, 1930, Verdooorn s.n. (holotype, 
FH) —cf. Verdoorn (1933). 


Dioicous. Plants small, up to 1.5 cm long, 0.5-0.7 mm wide, brown to dark 
brown in dried condition. Branching Lejeunea-type. Stem diameter 100- 
130 um Leaves imbricate, convolute when dry, widely spreading. Lobe 
oblong-ovate, 430-900 um long, 250-390 um wide, lateral margin irregularly 
toothed toward the apex, apex acute; cells of lobe thick-walled, light yellow, 
rhomboidal, trigones cordate, intermediate thickening frequent; marginal 
cells 8-13 x 5-7 um, mid-leaf cells 13-29 x 13-17 um, basal cells 16-38 x 6- 


Lejeuneaceae subfamily Ptychanthoideae 93 


14 um; oil bodies not seen. Lobule ovate, 150-270 um long, 50-90 um wide, 
inflated, apex obliquely truncate with small tooth, Underleaves closely 
imbricate, oblong, 200-490 um long, 150-650 um wide, margin entire, apex 
truncate to emarginate, toothed, insertion line shallowly curved. Androecia 
not seen. Gynoecia with 2 lejeuneoid innovations, bract ovate, 970-1380 um 
long, 440-550 um wide, toothed at apex and upper margin, apex acute, sinus 
up to 1/2 of lobe length; bracteole oblong to broadly oblong about as long 
as bracts or shorter, 500-670 um long, 350-410 um wide, margin serrulate, 
coarsely serrate towards the apex, apex emarginate. Perianth, oblong, 1800 
um long, 500 um wide, with 3 keels, margins serrulate in the upper part. 
Sporophytes and asexual reproduction not seen. 


Figure 24. Thysananthus minor Verd. Shoot (a); leaf lobe (b); underleaf (c); lobule (d); cells of 
midleaf (e); cells of basal part of the lobe (f). Drawn from Neervoort 1018, BO. 


Specimens examined: INDONESIA. West Java, Gn. Gede, above Cibodas, 
Geger Bentang, ca 1520-1700 m, Neervoort 1112, 1018, 1088, Noerta & 
Soekar 1139 (BO!); G. Halimun Nat. Park, ca 1000 m, Haerida 712 (BO!) 
ibid., ca 1200 m, collector unknown (BIOT!); Gn. Cikurai, W slope, ca 1700 
m Verdoorn 59g, 59h (BO!). 


Distribution: Java, Sumatra, Papua New Guinea. 


94 Gard. Bull. Singapore 62 (1) 2010 


Notes: Diagnostic characters of Thysananthus minor are: 1) very small plant, 
less than | mm wide, 2) leaf lobe irregularly toothed towards the apex, 3) 
underleaves toothed towards the apex, 4) lobule with only | small tooth. 
Thysananthus minor is new to Java. This species grows at ca 1000-1710 m 
and is very similar to 7. spathulistipus but differs by the smaller size. Possibly 
it is just a form of the latter species. 


25. Thysananthus retusus (Reinw. et al.) B. Thiers & Gradst., Mem. N.Y. Bot. 
Gard. 52:67 (1989); Jungermannia retusa Reinw et al., Acta Phys.-Med. Acad. 
Caes. Leop. Carol. Nat. Cur. 12: 214 (1824); Ptychanthus retusus (Reinw. et 
al.) Nees var. a, in Gottsche et al., Syn. Hepat. 292 (1845). —Type: Indonesia. 
Java, Blume s.n. (holotype, STR; isotype, W) — cf. Gradstein et al. (2002). 
—Thysananthus planus Sande Lac., Ned. Kruidk. Arch. 3,4: 419 (1854).—Type: 
Indonesia. Java, Junghuhn s.n. (holotype, L; isotype, G) — cf. Gradstein et al. 
(2002). 


{ 200Hm 


Figure 25. Thysananthus retusus (Reinw. et al.) B. Thiers & Gradst. Shoot (a); underleaves (b); leaf 
lobe (c); leaf lobule (d); female bracteole (e); female bract (f); vitta (g); cells of the margin of the 
leaf (h); cells of midleaf (i); perianth (j). Drawn from Haerida 511, BO. 


Plants up to 2 cm long, 0.7-1 mm wide; pale green to brown in the dried 
condition. Branching Lejewnea-type. Stem diameter 100-120 um. Leaves 
imbricate, convolute when dry, widely spreading. Lobe ovate to oblong, 740- 
840 um long, 350-550 um wide, margin entire, apex obtuse to acuminate; 
cells of lobe thick-walled, hyaline to light yellow, quadrangular to hexagonal, 


Lejeuneaceae subfamily Ptychanthoideae 95 


trigones triangular, intermediate thickening scarce; marginal cells 6-8 x 
3-5.5 um, mid-leaf cells 8-15 x 7-10 um, basal cells 13-19 x 5-8 um, vittae 
consisting of 2-3 rows of rectangular cells, ending at 2/3 of the lobe length, 
21-51 x 10-17 um; oil bodies not seen. Lobule oblong-ovate 290-310 um 
long, 140-150 um wide, apex with | tooth consisting of 3-4 cells. Underleaves 
imbricate, orbicular, obdeltoid to subrectanglar sometimes recurved 
seemingly emarginate, 230-310 um long, 290-390 um wide, margins entire, 
apex rounded, sometimes with scattered small teeth towards the apex. 
Generative structures not seen. 


Specimens examined: INDONESIA. West Java, Bogor Bot.Garden, W. Meijer 
55dlI (BO!); Gn. Salak, ca 2000 m, Kurz s.n. (BO!); Gn. Gede-Pangerango 
Nat. Park, Bodogol, Haerida 851, 811 (BO!, GOET!); G. Pancar, ca 400 m, 
Schiffner 287 (BO!); Geger Bentang, ca 1620 m, Neervoort 1070 (BO!). 


Distribution: Java, Philippines, West Irian, Papua New Guinea, Australia, 
Pacific Islands. 


Notes: Diagnostic characters of Thysananthus retusus are: 1) small plant with 
flattened leaves, 2) leaf with a distinct vitta, and with isodiametric non-vitta 
cells, 3) apical tooth of leaf lobule 3-4 cells long. This species grows at ca 400- 
2500 m. By the vitta, the small, isodiametric leaf cells and the long, curved 
lobule tooth T: retusus is a very distinct species that cannot be confused with 
any other member of Ptychanthoideae. 


26. Thysananthus spathulistipus (Reinw. et al.) Lindenb., in Gottsche et al., 
Syn. Hepat. 287 (1845); Jungermannia spathulistipa Reinw. et al., Acta Phys.- 
Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 212 (1824). -Type: Indonesia. 
Java, Bantam, Leback Mts., Blume s.n. (holotype, STR; isotypes, G, W) — cf. 
Gradstein et al. (2002). 


Autoicous. Plants robust, up to 3 cm long, 0.5-1 mm wide, brownish green to 
dark brown in dried condition. Branching Lejeunea-type. Stem diameter 80- 
180 um. Leaves imbricate, widely spreading. Lobe ovate-oblong, 820-1100 
um long, 320-630 um wide, margin coarsely toothed towards the apex, apex 
acute; cells of lobe thick-walled, light yellow, rhomboidal, trigones cordate, 
intermediate thickening frequent, cell walls at the basal part sometimes with 
orange color; marginal cells 10-20 x 7-10 um, mid-leaf cells 26-31 x 10-20 
um, basal cells 26-41 x 10-15 um; oil bodies not seen. Lobule ovate, 240-330 
um long, 40-90 um wide, inflated, apex truncate with | small tooth consisting 
of 1 cell. Underleaves imbricate, spathulate, 310-910 um long, 350-480 um 
wide, margin coarsely toothed towards the apex, apex truncate to sometimes 


96 Gard. Bull. Singapore 62 (1) 2010 


emarginate, insertion line almost straight. Androecia terminal on branches, 
bracts in 4-15 pairs, similar to leaves but lobules larger, bract lobule 2/3 of 
the bract lobe, bract lobe 270-370 um long, 150-160 um wide, hypostatic, 
bracteoles similar to underleaves in size and shape, 170-180 um long, 200-230 
um wide. Gynoecia on short or long branches, with 1-2 lejeuneoid innovations, 
bract ovate 900-1000 um long, 300-400 tm wide, crenulate towards the apex, 
apex acute, sinus up to 1/2 of lobe length; bracteole oblong to sub obovate 
700-2200 um long, 250-650 um wide, margin crenate, apex truncate. Perianth 
oblong, about 2000 um long, 550 um wide, with 3 keels, margins toothed. 
Sporophytes and asexual reproduction not seen. 


jb 


Figure 26. Thysananthus spathulistipus (Reinw. et al.) Lindenb. Shoot (a); leaf lobe (b); underleaf 
(c); female bracteole (d); female bract (e); cells of midleaf (f). Drawn from Haerida 725, BO. 


Specimens examined: INDONESIA. West Java, Bogor Bot. Garden, ca 250 
m, Schiffner 281 (BO!); Cibodas Bot. Garden, Verdoorn 20a, 20k, 301, 30p, 
300, Neervoort 1506, 70 (BO!); Gn. Pangrango, Meijer B895, B775, B286, 
B530, B447e, B519, B3339, B818c, B773a, B771a, B805, B81 1c, B81 1e, B385c, 
B752b, B753, B533c, B287b, B298b, B801a (BO!); Gn. Gede, 1500-1900 
m Verdoorn 46m, 46l, 46], 46h, 46g, 46f, Meijer B4093, Schiffner 291la, 289 
(BO!); Gn. Gegerbentang, E slopes, 1500-2000 m, Verdoorn 67f 67e, 67g, 
Neervoort 1139, 1144, 1985, 1022, 1014, Noerta & Soekar 1144, 50/1089, 
Meijer B637a (BO!); Gn. Halimun Nat. Park, Gradstein s.n., Haerida 725 
(BO!); Mt. Kendeng, ca 1060-1350 m, Radhiah Zakaria 173c, 181c, 218c, 
226c (BIOT!); G. Malabar Verdoorn 61g, 61f, 6le, 61d (BO!); Gn. Guntur, 
“Kawah Kamodjan”, 1500-1700 m, Verdoorn 50f, 50e (BO!); Gn. Cikurai, ca 
1700 m, Verdoorn 591 (BO!). 


Lejeuneaceae subfamily Ptychanthoideae 97 


Distribution: Java, Sumatra, Peninsular Malaysia, Borneo, Bali, Soembawa, 
Sulawesi, Moluccas, West Irian, Papua New Guinea, Australia, Solomon Is., 
Thailand, India, Sri Lanka, tropical Africa. 


Notes: Diagnostic characters of Thysananthus spathulistipus are: 1) 
leaves and underleaves symmetrical, with toothed margins, 2) spathulate 
underleaves, 3) coarsely toothed female bracts and bracteoles. This species 
grows at ca 200-2000 m and is most common in the mountains. It differs 
from other Javanese species of Thysananthus by the rather large plant size, 
the symmetrical, toothed leaves without vitta, the lobule with a short, blunt 
tooth, and the often narrowly spathulate underleaves. 


Discussion and conclusion 


This study revealed the occurrence of 26 species of Lejeuneaceae subfam. 
Ptychanthoideae in West Java, in 8 genera: Acrolejeunea (Spruce) Schiffn. 
(3 species), Archilejeunea (Spruce) Schiffn. (1 species), Lopholejeunea 
(Spruce) Schiffn. (10 species), Mastigolejeunea (Spruce) Schiffn. (3 species), 
Ptychanthus Nees (1 species), Schiffneriolejeunea Verd. (2 species), 
Spruceanthus Verd. (2 species) and Thysananthus Lindenb. (4 species). 
Two species found in this study were new records of the Hepaticae in 
Java: Mastigolejeunea indica and Thysananthus minor. The record of 
Mastigolejeunea indica is based on a specimen collected by the Dutch 
Bryologist, W. Meijer,in Bogor Botanical Garden in 1951 and in Meru Betiri 
National Park, East Java, in 2005. Five additional species recorded from Java 
in the literature (Acrolejeunea tjibodensis Verd., Dendrolejeunea fruticosa 
(Lindenb. & Gottsche) Lacout., Phaeolejeunea latistipula (Schiffn.) Mizut. 
[doubtful record], Spruceanthus sulcatus (Nees) Gradst. and Thysananthus 
comosus Lindenb.) were not found in this study. 

Based on the number of known localities in West Java the species 
of Ptychanthoideae can divided into several categories. Species with “wide 
distribution” are known from more than 5 localities, with “moderately wide 
distribution” from about 4, with “rather limited distribution” from about 
3, with “limited distribution” from about 2, and “very limited distribution” 
from only one locality. 

Lopholejeunea eulopha, Lopholejeunea ceylanica, Lopholejeunea 
subfusca and Thysananthus spathulistipus are the most common species in 
West Java, with a wide distribution. Thysananthus spathulistipus (200-1700 
m) was found in Bogor Bot. Garden, Cibodas, Gede Pangrango Nat. Park 
and Halimun Salak Nat. Park, and on Mt. Guntur, Mt. Cikurai and Mt. 
Malabar; Lopholejeunea eulopha (200-1700 m) in Bogor Bot. Garden, Ujung 


98 Gard. Bull. Singapore 62 (1) 2010 


Kulon Nat. Park and Halimun Salak Nat. Park, and on Mt. Guntur and Mt. 
Megamendung; L. ceylanica (500-1700 m) in Bogor Bot. Garden, Cibodas, 
Gede Pangrango National Park, Telaga Warna and on Mt. Cikurai; and L. 
subfusca (500-2500 m) in Bogor Bot. Garden, Cibodas, Gede Pangrango 
National Park and on Mt. Cikurai and Mt. Papandayan. 

Lopholejeunea nigricans, Lopholejeunea horticola and Thysananthus 
retusus have a moderately wide distribution. Lopholejeunea nigricans (200- 
1700 m) has been recorded from Bogor Bot. Garden, Mt. Pangrango, Mt. 
Guntur and Telaga Warna (Puncak Pass); L. horticola (800-2400 m) from 
Mt. Gede (including Cibodas), Mt. Guntur, Mt. Patuha and Mt. Malabar; 
and Thysananthus retusus (400-2500 m) from Bogor Bot. Garden, Mt. Gede, 
Mt. Halimun Salak Nat. Park and Mt. Pancar. 

Archilejeunea planiuscula, Lopholejeunea herzogiana, L. wiltensii, 
Mastigolejeunea virens, Ptychanthus striatus, Spruceanthus semirepandus, 
Thysananthus convolutus and Thysananthus minor have a rather limited 
distribution in West Java. Archilejeunea planiuscula (200-1450 m) was found 
- in Ujung Kulon Nat. Park and on Mt. Gede; Lopholejeunea herzogiana 
(1200-1500 m) in Bogor Bot. Garden, Gede Pangrango Nat. Park and Telaga 
Warna (Puncak Pass); L. wiltensii (1200-1900 m) in Gede Pangrango Nat. 
Park and on Mt. Guntur; Mastigolejeunea virens (200-1500 m) in Bogor 
Bot. Garden and on Mt. Gede and Mt. Megamendung; Ptychanthus striatus 
(1000-2400 m) in Gede Pangrango Nat. Park and on Mt. Megamendung; 
Spruceanthus semirepandus (1200-2300 m) on Mt. Gede, Mt. Cikurai and 
Mt. Malabar; Thysananthus convolutus (1300-2300 m) on Mt. Gede, Telaga 
Warna (Puncak Pass) and Mt. Malabar; and T. minor (1000-1700 m) on Mt. 
Gede, in Mt. Halimun Salak Nat. Park and on Mt. Cikurai. 

Acrolejeunea fertilis, A. pycnoclada, Lopholejeunea_ applanata, 
Mastigolejeunea auriculata and Schiffneriolejeunea tumida var. haskarliana 
were found in an even more limited distribution. Acrolejeunea fertilis was 
only found in the lowlands (60-80 m) in Pulau Panaitan and Depok; A. 
pycnoclada only at Mt. Gede and Ciater (Subang), in rather high altitude 
(1500-1900 m) although the species has also been recorded from lowlands 
(e.g. Bogor Bot. Garden) in the literature; Schiffneriolejeunea tumida var. 
haskarliana (440-1500 m) only from Mt. Gede and Mt. Halimun Salak Nat. 
Park; Lopholejeunea applanata (ca 1200 m) only from Gede Pangrango 
Nat. Park; and Mastigolejeunea auriculata (200-500 m) only from Bogor 
Bot. Garden and Ujung Nat. Park. This condition may be due to inadequate 
collections from the area. 

Acrolejeunea arcuata, Lopholejeunea recurvata, L. zollingeri, Mastigolejeunea 
indica, and Schiffneriolejeunea pulopenangensis, finally, had the most limited 
distribution and were found in only one location, i.e. Acrolejeunea arcuata 
only on Mt. Patuha at 2000 m, Lopholejeunea recurvata on Mt. Gede at ca 


Lejeuneaceae subfamily Prychanthoideae 99 


1450 m, L. zollingeri on Mt. Pangrango at ca 1600 m, and Mastigolejeunea 
indica and Schiffneriolejeunea pulopenangensis only in Bogor Bot. Garden 
at about 200 m. The records of the latter two species only from Bogor 
Botanical Garden emphasize the importance of this garden as a habitat for 
Ptychanthoideae in West Java. A searching for additional localities of the 
rare species in West Java is needed. 

According to Gradstein (1991) the endemic genera of Asiatic 
Ptychanthoideae are largely restricted to the subtropical and temperate 
areas of Asia and some are also known as fossils in Eocenic amber of Europe. 
They are considered to be palaeoendemic, the relictual groups. Endemic 
genera of Asiatic Lejeuneoideae, however, occur mainly in the tropical rain 
forests of the Malesian archipelago, are often highly specialized, and are 
lacking in the fossil record. They probably have co-evolved in the Tertiary 
with the rain forest and are to be considered neoendemics. 

Ptychanthoideae seem to be older than Lejeuneoideae and may 
already have existed in the Mesozoic before the break-up of Laurasia and 
Gondwanaland. The Mesozoic age was recently confirmed based on fossil 
evidence and DNA sequence analysis by Wilson er al. (2007), who found 
that Lejeuneaceae started to diversify in the Late Cretaceous, about 60-90 
million years ago. 

The geographical distribution and altitudinal ranges of Ptychanthoideae 
in West Java are shown in Table 1; the definition of the distribution types 
follows Ariyanti and Gradstein (2007). The geographical ranges of the 
species were determined based on collected specimens and literature (e.g., 
Mizutani, 1961; Gradstein and Terken, 1981; Menzel 1988; Gradstein er al., 
2002; Zhu and Gradstein, 2005: Aniyanti and Gradstein, 2007). 

The data show that the species of Ptychanthoideae of West Java can be 
subdivided into 4 groups by their geographical distributions: Malesian species 
(8 spp.), tropical Asiatic species (10 spp.), palaeotropical species (tropical 
Asia + Africa; 3 spp.) and pantropical species (throughoput the tropics: 4 
spp.). It appears that the species are rather widespread; none of the species 
are endemic to Java or western Malesia. The widespread distribution of the 
species is probably due to their dispersal by spores, which may be easily 
carried by the wind over long distances (van Zanten and Gradstein, 1987). 
But also the rather old age of Ptychanthoideae (Gradstein, 1991: Wilson er 
al., 2007) may play a role. 

The majority of the species (about 20) are found at mid-montane 
elevations, at 1200-1500 m. Few species found below 100 m and above 2000 
m. The data from West Java agree with the general pattern of altitudinal 
distribution reported for Lejeuneaceae. According to Gradstein (1995), 
the diversity of Lejeuneaceae decreases with elevation and accounts 
for about 45% of total hepaticae diversity in the lower montane forest 


100 Gard. Bull. Singapore 62 (1) 2010 


Table 1. Altitudinal and geographical distributions of the species of Ptychanthoideae 
recorded in West Java. Lowland: 0-1200 m. Montane: 1200-3000 m. A: Asiatic. M: Malesian. P: 


Pantropical. Pal: Palaeotropical (Asia, Africa). * species new to Java. 


Altitudinal — distribution Geographical 


Nip Species Lowland Montane distribution 
1. | Acrolejeunea arcuata - + M 
2. Acrolejeunea fertilis + - M 
3 Acrolejeunea pycnoclada + + Pal 
4 Archilejeunea planiuscula ~ + A 
> Lopholejeunea applanata - . A 
6. | _Lopholejeunea eulopha + + P 
7 Lopholejeunea herzogiana - Ta M 
8 Lopholejeunea nigricans + + P 
J) Lopholejeunea ceylanica + + A 
10. Lopholejeunea horticola + a A 
11. Lopholejeunea recurvata + + M 
12. Lopholejeunea subfusca + + 2 
13. Lopholejeunea wiltensii - + M 
14. Lopholejeunea zollingeri - + A 
15. Mastigolejeunea auriculata + + Je 
16. | Mastigolejeunea indica* + - M 
17. Mastigolejeunea virens + + M 
18. Ptychanthus striatus +: + Pal 
19. Schiffneriolejeunea 

pulopenangnesis oF : M 

20. — Schiffneriolejeunea tumida var. 

haskarliana - a A 

21.  Spruceanthus polymorphus + + A 

22.  Spruceanthus semirepandus - + A 

23. Thysananthus convolutus - + A 

24. = Thysananthus minor* + + M 

25.  Thysananthus retusus + + A 


u 
= 


26. Thysananthus spathulistipus + + 


Lejeuneaceae subfamily Prichanthoideae 101 


(1000/1400-2000/2500 m). 30% in the upper montane forest (2000/2500- 
3000/4000 m) and 20% in the subalpine forest (above 3000/4000 m). 

The members of the Ptychanthoideae of West Java are epiphytes and 
grow on the bark of trees. on treelets and shrubs. As indicated by Thiers 
and Gradstein (1989) and Gradstein et al. (2001). many species are rather 
xerotolerant epiphytes of trees at forest margins or in rather open vegetation. 
None of the species of West Java were found growing on living leaves. which 
are generally inhabited by tiny members of the subfamily Lejeuneoideae. 
and few species occur on small branches of shrubs. which are the habitat 
of the ramicolous bryophytes. Dendroid. feather or bracket-type mosses 
and liverworis. belonging to Neckeraceae, Hookeriaceae, Pterobryaceae. 
and Plagiochilaceae. as well as various tiny members of Lejeuneaceae are 
the specialists of this habitat (Gradstein and Pocs, 1989). A study of the 
ecology and optimum habitat conditions of the members of Lejeuneaceae 
subfam. Ptychanthoideae, including moisture and temperature rates, 
would be desirable. Such data might further improve our understanding 
of the distribution of the species of Ptychanthoideae in West Java. 


Acknowledgements 


The first author is grateful to the ~Program Karyasiswa Dalam Negeri LIPI” 
for financial support of her study. She also thanks Dr. Johanis Palar Mogea 
for his valuable comments on the manuscript. Furthermore. she gratefully 
acknowledges the support of the Directors of Herbarium Bogoriense and 
Herbarium Biotrop for providing facilities to conduct her research. 


References 


Anriyanti, N.S. and S.R. Gradstein. 2007. Wallace’s line and the distribution of 
the liverworts of Sulawesi. Cryptogamie. Bryologie 28: 3-14. 

Gradstein, S.R. 1975. Monograph of the genus Acrolejeunea. Bryophytorum 
Bibliotheca 4- 1-216. 


Gradstein, S.R. 1991. Diversity and distribution of Asian Lejeuneaceae 
subfamily Ptychanthoideae. Tropical Bryology 4: 1-16. 


Gradstein. S.R. 1994. Verdoorm’s Studien iiber Asiatische Jubulae. Hikobia 11: 
451-456. 


102 Gard. Bull. Singapore 62 (1) 2010 


Gradstein, S.R. 1995. Diversity of Hepaticae and Anthocerotae in montane 
forests of the tropical Andes, pp. 321-334. In: Churchill, S.P., J. Luteyn, E. 
Forero and H. Balslev (eds.), Biodiversity and Conservation of Neotropical 
Montane Forests. New York Botanical Garden, New York. 


Gradstein, S.R., S.P. Churchill and N. Salazar-Allen. 2001. Guide to the 
bryophytes of Tropical America. Memoirs of the New York Botanical 
Garden 86: 1-157. 


Gradstein, S.R., X.-L. He, S. Piippo and M. Mizutani. 2002. Bryophyte flora 
of the Huon Peninsula, Papua New Guinea. LX VIII. Lejeuneaceae 
subfamily Ptychanthoideae (Hepaticae). Acta Botanica Fennica 174: 1- 
88. 


Gradstein, S.R. and T. Pocs. 1989. Bryophytes, pp. 311-325. In: Lieth, H. and 
M.J.A. Werger (eds.), Tropical Rain Forest Ecosystems. Elsevier Science 
Publishers, Amsterdam. 


Gradstein, S.R. and L.Terken. 1981. Studies on Lejeuneaceae subfam. 
Ptychanthoideae VI. A revision of Schiffneriolejeunea sect. Saccatae 
from Asia. Occasional Papers of the Farlow Herbarium of Cryptogamic 
Botany 16: 71-81. 


Hasan, M. and N.S.Ariyanti. 2004. Mengenal Bryophyta (Lumut) Taman 
Nasional Gunung Gede Pangrango. Vol. 1. BalaiTaman Nasional Gunung 
Gede Pangrango, 93 pp. 


Menzel, M. 1988. Annotated Catalogue of the Hepaticae and Anthocerotae 
of Borneo. Journal of the Hattori Botanical Laboratory 65: 145-206. 


Mizutani, M. 1961. A revision of Japanese Lejeuneaceae. Journal of the 
Hattori Botanical Laboratory 24: 116-180. 


Mizutani, M. 1969. Lejeuneaceae subfamily Ptychanthoideae from Sabah 
(North Borneo). Journal of the Hattori Botanical Laboratory 32: 129- 
139: 


Nees von Esenbeck, C.G. 1830. Enumeratio Plantarum Cryptogamicarum 
Javae et Insularem Adjacentium. I. Hepaticas complectens. Breslau. 


Piippo, S, T. Koponen and D.H. Norris. 1987. Endemism in the Bryophyte 
Flora in New Guinea. Symposia Biologica Hungarica 35: 361-372. 


Lejeuneaceae subfamily Ptychanthoideae 103 


Reinwardt, C., C. Blume and C.G. Nees von Esenbeck. 1824. Hepaticae 
javanicae. Acta Physico-Medica Academiae Caesareae Leopoldino- 
Carolinae Naturae Curiosorum 12: 181-238. 


Sande Lacoste, C.M. 1856. Synopsis Hepaticarum Javae. Verhandelingen der 
Koninklijke Nederlandse Akademie van Wetenschappen 5: 1-112. 


Schiffner, V. 1898. Conspectus Hepaticarum Archipelagi Indici. Batavia 
Staatsdruckerei. 382 pp. 


Schuster, R.M. 1980. Phylogenetic studies on Jungermanniidae IT. Radulineae 
(Part I). Nova Hedwigia 32: 637-693. 


Thiers, B.M. and S.R. Gradstein. 1989. Lejeuneaceae (Hepaticae) of 
Australia. I. Subfamily Ptychanthoideae. Memoirs of the New York 
Botanical Garden 52: 1-79. 


Verdoorn, F. 1933. Die von V. Schiffner (1893-1894) und von Fr. Verdoorn 
(1930) auf den Indomalesischen Inseln gesammelten Lejeuneaceae 
Holostipae. De Frullaniaceis XI. Recueil des Travaux Botaniques 
Néerlandais 30: 212-233. 


Verdoorn, F. 1934. Studien tiber Asiatische Jubuleae. Annales Bryologici 
Supplement 4: 1-231. 


Wilson, R., J. Heinrichs, J. Hentschel, S.R. Gradstein and H. Schneider. 2007. 
Steady diversification of derived liverworts under steady Tertiary climatic 
fluctuations. Biology Letters 3: 566-569. 


Zanten,B.O.van,andS.R.Gradstein. 1987.Experimental dispersal geography 
of tropical liverworts. Nova Hedwigia Beiheft 90: 41-94. 


Zhu, R.-L. and S.R. Gradstein. 2005. Monograph of Lopholejeunea 
(Lejeuneaceae, Hepaticae) in Asia. Systematic Botany Monographs TA: 
1-98. 


2 ass - 


Gardens’ Bulletin Singapore 62 (1): 105-109. 2010 105 


Phaius takeoi (Orchidaceae) Newly Recorded from 
Thailand and Myanmar 


H. KURZWEIL .S.WATTHANA AND S.LWIN®> 


Herbarium, Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569 
Queen Sirikit Botanic Garden, The Botanical Garden Organization. 
, P.O. Box 7, Mae Rim, Chiang Mai 50180, Thailand 
~ Myanmar Floriculturist Association, Alone Road, Yangon, Myanmar 


Abstract 


The occurrence of Phaius takeoi (Hayata) H.J. Su (Orchidaceae), previously 
only known in mainland China, Vietnam and Taiwan, is here reported for 
Thailand and Myanmar. 


Introduction 


In the course of floristic and taxonomic work on the orchids of Thailand and 
Myanmar three collections of a yellow and green-flowered Phaius species 
were made which did not match any currently known species (Seidenfaden, 
1986; Kress et al.,2003; Vaddhanaphuti, 2005, Kurzweil, 2010 ). The specimens 
are now positively identified as P. takeoi (Hayata) H.J. Su, currently only 
known in mainland China, Vietnam and Taiwan. The new record of this 
species in Thailand and Myanmar is reported below. 


Phaius takeoi (Hayata) H.J. Su 

Quart. J. Exp. Forest Natl. Taiwan Univ. 3 (1989) 77; Chen er al., Fl. China 
vol. 25: 290 (2009): Su, in Digital Flora of Taiwan, http://www.efloras.org 
florataxon.aspx/flora_id =100&taxon_id =242413999, accessed January 12, 
2010). —Basionym: Calanthe takeoi Hayata, Icon. Pl. Formosan. 9 (1920) 111. 
—Typus: Taiwan, Toyencho, Urai, Oct 1917, Takeo Ito s.n. (not found). Plate 1. 
—Phaius longicruris Z.H.Tsi, Acta Phytotax. Sin. 19 (1981) 505; Averyanov 
and Averyanova, Turczaninowia 5 (2002) 93. —Typus: China, Yunnan, 
Jinghong, in forest, 1400 m alt., Oct 1936, Wang 79184 (holotype, PE). 


Terrestrial herbs, vegetative parts entirely glabrous, basally pseudobulbous, to 
1 m tall. Roots to 3 mm in diameter. Pseudobulbs elongate-clavate, 20-40 cm 
long and (1.3-)2-2.5 cm in diameter, slightly tapering upwards, with 3-5 nodes, 
internodes 2.5-10.7 cm long, green or dark green with green veins, glabrous, 


106 Gard. Bull. Singapore 62 (1) 2010 


Plate 1. Phaius takeoi (Hayata) H.J. Su. A. Plant; B. Part of inflorescence; C & D. Flowers. All 
from Watthana 3287 (Thailand, Tak); D flower photo from Nyan Tun s.n. (Myanmar, Shan 
State). Bars: A. 10 cm. B-D.3 cm. 


Phaius takeoi (Orchidaceae) 107 


covered by pale green sheaths. Leaves 5-8, arising from the apical portion of 
the pseudobulbs, elliptic, elliptic-lanceolate or elliptic-oblong. (5.5-)12-40 x 
(2.5-)5.5-10(-14) cm, acuminate or long-acuminate, margin entire and slightly 
undulate, with 5-7 veins, green above, pale green underneath. Inflorescences 
arising from the lower nodes of the pseudobulbs, erect. up to 60 cm long 
and 0.4-0.8 cm in diameter, normally racemose but once observed with a 
side branch (in the specimen Watthana 3287), glabrous below, glabrous or 
sparsely hairy on the uppermost part of the rachis. Peduncles 30-50 cm long, 
with up to 9 sterile bracts 16-28 mm long. Rachis 10-13 cm long (rachis in all 
three examined specimens still elongating), with 4-15 flowers. Floral bracts 
persistent, ovate-lanceolate or narrowly lanceolate, the lower ones to 17 x 
3.8 mm, acute, glabrous. Flowers 4-5 cm in diameter, opening widely: sepals 
and petals pale green or yellowish green and sometimes darker at the apex. 
lip white or cream and sometimes yellow-tinged. Pedicel plus ovary 2-3.2 cm 
long. sparsely pubescent or nearly glabrous. Sepals subequal. elliptic-oblong. 
obtuse or acute, 27-32(-35) x (7-)8-10.5 mm, 5-7-veined, abaxially sparsely 
hairy: laterals slightly oblique, somewhat decurved. Petals oblanceolate or 
oblanceolate-oblong, sometimes basally indistinctly clawed, obtuse or acute. 
28-33 x 6-8 mm, 3-veined. Lip nearly orbicular, 30-34 x 24-26 mm (excluding 
spur), shallowly 3-lobed, glabrous for the most part but basal portion and 
inside of spur hairy, basally united with the column for 7-8 mm: midlobe 
orbicular-square or suborbicular, apex emarginate and sinus sometimes 
mucronate, margins undulate: side lobes ovate or elongate-ovate, strongly 
incurved and forming a wide open tube to embrace the column, apex broadly 
rounded, margins undulate: disc with three faint yellow and smooth ridges, 
the median one extending from the base of the lip to just below the apex of 
the midlobe, the lateral ones arising from the middle part of the lip to the 
base of the midlobe, spur elongate-conical with broad entrance, clavate, 6- 
9 mm long, greenish yellow. Column stout, 15-25 mm long, 6-7 mm broad, 
white, hairy on the front face below the stigma and continually into the spur. 
apex widened with very prominent lateral appendages. 


Specimens examined: THAILAND. Northern Region. Phitsanulok Province. 
Phu Soi Dao, Sep 2008, RMK 671/51 (QBG, QBG spirit): Northern Region. 
Tak Province, Umphang, Jul 2009, Watthana 3287 (QBG). MYANMAR. 
Shan State, Taunggyi, Jul to Aug 2009, Nyan Tun s.n. (SING). 


Habitat and flowering time: This species is found in moist, broad-leaved 
primary or secondary forest. The Thai specimens were found in a marshy 
spot in evergreen forest, while the Myanmar specimen was collected in moist 
soil near streams. Populations were quite sizeable in the Myanmar plant 
where several dozen individuals were found on a hill slope. but very small 


108 Gard. Bull. Singapore 62 (1) 2010 


with under 20 individuals in the two Thai specimens’. Altitudes were not 
recorded in the Thai and Myanmar specimens reported here, but elsewhere 
the species is found at 500-1400 m (Averyanov and Averyanova, 2002; Tsi, 
1981: Chen et al.,2009; Su, no date, Digital Flora of Taiwan). Flowering occurs 
between July and September, and has elsewhere been reported between 
October and December. In Taiwan and Vietnam the species is said to be 
uncommon (Averyanovy and Averyanova, 2002; Su, no date, Digital Flora of 
Taiwan). 


Distribution: China (Yunnan), Taiwan, Vietnam, Thailand, Myanmar. 


Notes: Among the Thai and Myanmar Phaius congeners, this species is 
unmistakable with its habit and the yellow-green flowers. 


Acknowledgements 


- We would like to acknowledge the Queen Sirikit Botanic Garden for 
assistance during fieldwork and the Rom Klao Botanical Garden for 
providing one of the specimens examined. We would also like to thank 
the collector of the Myanmar specimen, Mr. Nyan Tun, for providing 
information, and the Myanmar Forest Department for issuing a CITES 
permit to transport the plant to Singapore for identification. Dr. Jin Xiaohua 
is thanked for information on the type specimen of Phaius longicruris. 


References 


Averyanov,L.andA.Averyanova.2002.Rare species of orchids (Orchidaceae) 
in the flora of Vietnam. Turczaninowia 5: 49-108. 


Chen S-C, Liu Z-J, Zhu G-H, Lang K-Y,Tsi Z-H, Luo Y-B, Jin X-H, P.J. Cribb, 
J.J. Wood, S.W. Gale, P. Ormerod, J.J.Vermeulen, H.P. Wood, D. Clayton 
and A. Bell. 2009. Orchidaceae. In: Wu. Z-Y., P-H. Raven and Hong. D.- 
Y. (eds.), Flora of China, vol. 25, pp. 1-566. Science Press, Beijing, and 
Missouri Botanical Garden Press, St. Louis. 


Kress, W.J., R. DeFilipps, E. Farr and Yin-Yin-Kyi. 2003. A checklist of the 
trees, shrubs, herbs and climbers of Myanmar. Contributions from the US 
National Herbarium 45: 1-590. 


Kurzweil, H. 2010. A precursory study of the Calanthe group (Orchidaceae) 
in Thailand. Adansonia, sér. 3, 32: 57-107 


Phaius takeoi (Orchidaceae) 10° 


Su. H-J. Not dated. Phaius. In: Digital Flora of Taiwan. Published on the 
internet at http://www.efloras org/florataxon.aspx/flora_id=100&taxon_ 
id=242413999_ accessed January 12.2010. 


Seidenfaden, G. 1986. Orchid genera in Thailand XIII. Thirty-three 
epidendroid genera. Opera Botanica 89: 1-216. 


Tsi, Z-H. 1981. New species of Orchidaceae from China. Acta 
Phytotaxonomica Sinica 19: 505-510. 


Vaddhanaphuti, N. 2005. A field guide to the wild orchids of Thailand, ed. 4. 
Silkworm Books. Chiang Mai. 


5 =z 


Gardens’ Bulletin Singapore 62 (1): 111-117. 2010 Wl 


Curcuma vitellina (Zingiberaceae), a New Species 
from Vietnam 


J. LEONG-SKORNICKOVA |, TRAN H.D.” AND M.E NEWMAN’: 


‘The Herbarium, Singapore Botanic Gardens 
5 1 Cluny Road, 259569 Singapore 
University of Science, Vietnam National University 
227 Nguyen Van Cu, QO5, Ho Chi Minh City, Vietnam 
* Royal Botanic Garden Edinburgh 
20A Inverleith Row, Edinburgh EH3 5LR, Scotland. 
Author for correspondence: jana_skornickova@seznam.cz 


Abstract 


Curcuma vitellina,anew species of Zingiberaceae from Tay Nguyén, Vietnam, 
is described, illustrated and compared to its closest ally, C. pierreana. 


Introduction 


Cambodia, Laos and Vietnam reportedly form a diversity hotspot for the 
family Zingiberaceae, although detailed data are lacking because the most 
recent comprehensive account of the family there is over a century old 
(Gagnepain, 1908). The main centres of diversity of the genus Curcuma 
are usually said to be India, Burma and Thailand (e.g. Leong-Skorniékova 
et al. 2008), but our recent explorations of Zingiberaceae for the Flora of 
Cambodia, Laos and Vietnam indicate that these countries are at least as 
rich in Curcuma and other Zingiberaceae. 

Three Curcumaspecies, C. bicolor Mood & K.Larsen,C. glans K. Larsen 
& Mood from Thailand and C. rhomba Mood & K. Larsen from Vietnam 
were described in 2001 (Mood & Larsen 2001). As a revision of Curcuma in 
Cambodia, Laos and Vietnam progresses,specimens collected at two localities 
in Vietnam have been shown to represent a new species which is described 
and illustrated here. Like the species described by Mood and Larsen, this 
new one lacks a clear distinction between fertile and coma bracts, but differs 
in overall shape and coloration of the flower as well as anther morphology. 


Curcuma vitellina Skorniék. & H. D. Tran, sp. nov. 

Curcumae pierreanae comae absentia, bractearum formae, anthera basi 
calcaribus filamentaceis 2 similis, sed rhizomate ramis lanceolatis verticaliter 
fasciculatis (contra rhizomate ramis horizontaliter repentibus), corollae lobis 


ee Gard. Bull. Singapore 62 (1) 2010 


dilute luteis (contra albis), labello staminodtisque lutee aurantiacis (contra 
albis vel albis apicibus profunde rosee purpurascentibus), lamina tenui 
valide plicata basi rotundata (contra coriacea vents principalibus minus 
prominentibus, basi cuneata ad attenuata) differt. -Typus: Vietnam, Lam 
Dong Prov., Pongour waterfall; 11° 41’ 07.0” N, 108° 16’ 06.1” E; 787 m; 
23 Jun 2008, Tran et al. 70 (holotype, SING incl. spirit; isotype, E, P, VNM, 
National University of Laos). 


Rhizome ovoid, ca 2.5-5 x 1.5-3 cm, with lanceolate to narrowly lanceolate 
branches held upright, 3-8 x 1-1.5 cm, brown externally, light yellow internally, 
slightly aromatic, root tubers elliptic, 2-4 cm long, light brown externally, 
cream white internally, at 5-15 cm from rhizome. Pseudostem to 15 cm long, 
green, composed of leaf sheaths and enclosed by 2 sheathing bracts, ligule to 
5 mm long, bilobed, hyaline, greenish white, translucent, turning papery with 
age, hairy at the apex, hairs ca 0.3 mm long; leafy shoot to 70 cm tall with up to 
5-6 leaves when flowering; petiole 5-20 cm long (petiole of first leaf shortest, 
innermost leaves longest), green, glabrous; lamina elliptic to elliptic-ovate, 
20-45 x 8-15 cm, glabrous on both surfaces, prominently plicate, adaxially 
bright green, shiny, abaxially lighter green, shiny; midrib glabrous, green; 
base rounded, margin hyaline, translucent white, ca 0.5 mm wide, glabrous; 
apex acute, shortly hairy. Inflorescence central, many flowered. Peduncle 4- 
20 cm long, up to 1 cm diam., greenish-white, puberulent, embedded within 
pseudostem. Spike 8-15 cm long, ca 4-5 cm diam. at the middle, without 
coma. Fertile bracts 15-60, larger at the base of the inflorescence, ca 3.5-4.5 
x 2.5-3.5 cm, ovate to trullate, smaller and ovate at the apex, cream white or 
pale greenish, sometimes with slight pinkish tinge, both sides shortly densely 
pubescent, connate in the lower 1/3 to 1/4. Cincinni with 4-6 flowers at the 
base of the inflorescence, 2-3 flowers at the top. Bracteoles one per flower, 
ovate, boat-shaped, ca 7 x 4 mm to 15 x 7 mm (outer ones larger, inner 
ones are gradually smaller), hyaline, translucent white, glabrous, but for the 
apex, upper part and margins sparsely hairy. Flowers 5-5.5 cm, exserted from 
bracts. Calyx ca 17 mm long, teeth 3, unilaterally split ca 7 mm, translucent 
white, sparsely hairy on the three main veins leading from the tooth to the 
base. Floral tube ca 3 cm long, narrowly cylindrical at base for ca 2 cm above 
the ovary, funnel-shaped at apex, externally white turning pale yellowish 
towards the apex, with glandular hair, internally white with dorsally placed 
groove holding the style: dorsal corolla lobe ca 20 x 11 mm, triangularly 
ovate, concave, glabrous, cream white with pale yellowish apex, apex 
mucronate, mucro less than 1 mm long with a few short hairs; lateral corolla 
lobes ca 18 x 9 mm, triangular with a rounded, slightly concave apex, cream 
white with pale yellowish apex, glabrous. Lateral staminodes obovate, ca 18 
x 11 mm, light yellow at base, yellow-orange towards the apex, glandular 


Curcuma vitellina (Zingiberaceae) from Vietnam 113 


hairs present on the raised middle portion facing the centre of the flower. 
Labellum ca 21 x 20 mm, obscurely trilobed, lateral lobes folding upwards, 
middle lobe emarginate with an incision up to 8 mm long, which splits at the 
apex of the lobe as flowering progresses, cream white at base, yellow at apex 
with deep yellow-orange band running through the centre (golden median 
band). Filament 4-6 mm long, pale yellowish, 4.5 mm at base, 2 mm at apex, 
with glandular hair at the back. Anther spurred, connective densely covered 
with short glandular hairs, anther spurs ca 0.5 mm long, filamentous, cream 
white, anther crest present, 1-1.5 x ca 2 mm, deep yellow, anther thecae 5 
mm long, white, dehiscing along their whole length. Style white, glabrous, 
stigma ca 1 mm wide, white, ostiole facing upwards, ciliate. Epigynous glands 
two, cream, 4 x 0.8 mm, with blunt apex. Ovary 3 x 2 mm, trilocular, hairy, 
hairs ca 0.2 mm long. Fruits not seen. 


Flowering: June to August. 


Distribution & habitat: So far known only from two localities (Bao Loc Pass 
and Pongour Waterfall) in Lam D’6ng province, Tay Nguyén, Vietnam. It 
grows among rocks in open and semi-open shrubby vegetation. 


Vernacular names & uses: None so far recorded. 


Etymology: Medieval Latin vitellinus, from Latin vitellus, egg yolk, referring 
to the deep yellow colour of the flowers. 


Other specimens examined: Among the vast amount of Curcuma herbarium 
material revised and digitised by the first author, we have found only one 
collection from Pongour (24 Aug 1924, Evrard 1193 x 2, P), which can be 
identified with certainty as C. vitellina. There are a few other specimens 
collected within a radius of 100 km of Pongour, which might represent C. 
vitellina (e.g. Poilane 30582, P and Poilane 5031 x 2, P), but the condition 
of the specimens and poor accompanying notes do not allow a definite 
identification to be made. 


IUCN preliminary assessment: Endangered (E): B2 ab (iii). The area of 
occupancy is estimated to be less than 500 km. The species is known to 
exist at only two locations with continuing decline in extent and quality of 
habitat, which is disturbed by erosion and development for tourism. 


114 Gard. Bull. Singapore 62 (1) 2010 


Figure 1. Curcuma vitellina, sp. nov. A. Habit; B. Inflorescence; C. Flower (front view): D. 
Detail of anther (front and side view); E. Liguie; F Rhizome and base of leafy shoot: G. 
Dissected flower. Photographs by J. Leong-Skorniékova taken from Tran et al. 70. 


Curcuma vitellina (Zingiberaceae) from Vietnam MNS 


Figure 2. Curcuma pierreana Gagnep. A. Habit; B. Inflorescence; C. Anther (front and side 
view) Photographs by J. Leong-Skorniékova from Tran et al. 26. 


Notes: It is expected that Curcuma vitellina will be a seed-setting species as 
the presence of young seedlings in natural populations has been observed. 
Curcuma vitellina is similar to C. pierreana by its inflorescence composed 
of cream or greenish flower bracts (which may be tinged pink), reflexed 
at the tips and lacking a distinct coma (the inflorescence of C. vitellina is 
more robust and with more bracts than that of C. pierreana). The anthers 
of both species have a well-developed crest and two filamentous spurs at 
the base, but the overall shape of the anther is different (see Figs. 1-3). The 
two species differ in their rhizomes, the branches of which are narrowly 


116 Gard. Bull. Singapore 62 (1) 2010 


Figure 3. Anthers in side, front, and back view. A. Curcuwma vitellina (Tran et al. 70); B. 
Curcuma pierreana (Tran et al. 26). Photographs by H.D. Tran. 


lanceolate and vertically clustered in C. vitellina, but creep horizontally in 
C. pierreana. The lamina in C. vitellina is thin, prominently plicate, glabrous 
on both sides and has a rounded base, whilst it is rather leathery with dense, 
short hairs abaxially, with less prominent venation, and with a cuneate to 
attenuate base in C. pierreana. The flowers are yellow to yellow-orange in 
C. vitellina, while C. pierreana has white flowers with a yellow band on the 
labellum and dark maroon tips to the labellum and lateral staminodes. 


Acknowledgments 
We thank the curators of BK, BKF, E, K, P, SING, VNM herbaria for letting 


us examine specimens in their care, the Asian Zingiberaceae Information 
Centre at Singapore Botanic Gardens, and the Zingiberaceae Resource 


Curcuma vitellina (Zingiberaceae) from Vietnam 117 


Centre at the Royal Botanic Garden Edinburgh (http://elmer.rbge.org.uk/ 
ZRC/) for providing protologues and related references. We are grateful to 
Prof. Lé Cong Kiét and Dr. Tran Triét for their support during our fieldwork 
in Vietnam, and to Dr. J.F. Veldkamp (L) for translating the diagnosis into 
Latin. The second author thanks Singapore Botanic Gardens for granting 
an SBG Fellowship to work on the Zingiberaceae of Cambodia, Laos 
and Vietnam. The fundings by Sud Expert Plantes, France [SEP project 
350], by the National Parks Board (Singapore), and by the Czech Science 
Foundation, GACR [grant numbers 521/09/0202 and P506/10/0623] are 
gratefully acknowledged. 


References 


Gagnepain, F. 1908. Zingibéracées, pp. 25-121. In: Lecomte, H. (ed.), Flore 
Générale de ’ Indo-Chine, vol. 6. Masson & Co., Paris. 


Leong-Skorniékova, J. , Sida, O., Sabu, M. & K. Marhold. 2008. Taxonomic 
and nomenclatural puzzles in Indian Curcuma: the identity and 
nomenclatural history of C. zedoaria (Christm.) Roscoe and C. zerumbet 
Roxb. Taxon 57: 949-962. 


Mood, J. & K. Larsen. 2001. New curcumas from South-east Asia. The New 
Plantsman 8: 207-217. 


NY 
4 


Gardens’ Bulletin Singapore 62 (I): 119-120. 2010 119 


New Combinations in Haplopteris ( Adiantaceae) for the 
Flora of Peninsular Malaysia 


S. LINDSAY 


Royal Botanic Garden Edinburgh, 
20A Inverleith Row 
Edinburgh EH3 5LR, Scotland, UK. 


Abstract 


Two new combinations are made here: Haplopteris hirta (Fée) S.Linds. and 
Haplopteris angustissima (Holttum) S.Linds. 


In preparation for the account of the Adiantaceae for the Flora of Peninsular 
Malaysia new combinations are necessary in the genus Haplopteris C.Presl. 
The family delimitation to be used in the Flora account is modified from 
Smith et al. (2006, 2008) who recognized a broadly delimited Pteridaceae 
but acknowledged that there are five monophyletic groups within the family 
that could be raised back to family level. The Adiantaceae, comprising 
Adiantum L. and all members of the former family Vittariaceae, is one of 
these monophyletic groups. 

As so defined, the Adiantaceae in Peninsular Malaysia consists of 
the five genera Adiantum L., Antrophyum Kaulf.. Haplopteris C.Presl, 
Monogramma Comm. ex Schkuhr., and Vaginularia Fée. The generic 
delimitation of the Vittarioid genera in the Flora account will largely 
follow Crane (1998). In this work it was shown that the Old and New 
World Vittaria species did not form a monophyletic group. All Vittaria 
species in Peninsular Malaysia (see Holttum, 1955) now belong in the 
genus Haplopteris. Crane (1998) and Zhang (2003) made most of the 
mecessary name changes but two new combinations are still required. 


Haplopteris hirta (Fée) S.Linds. comb. nov. 
Basionym: Vittaria hirta Fée, Mém. Foug. 10 (1865) 12. -Type: Wallace s.n., 
Borneo (holotype, BM). 


Haplopteris angustissima (Holttum) S.Linds.. comb. nov. 

Basionym: Vittaria angustissima Holttum, Gard. Bull. Singapore 11 (1947) 
274. -Type: C_E.Carr s.n., Pahang. Fraser’s Hill, alt. 4000 ft. March 
1929 (holotype, SING). 


120 Gard. Bull. Singapore 62 (1) 2010 


Acknowledgements 


I thank the Forest Research Institute Malaysia (FRIM) for a visiting 
Fellowship to pursue research on Malaysian Adiantaceae. I also thank 
Serena Lee (SING) and Alison Paul (BM) for their help in locating type 
material. 

References 


Crane, E.H. 1998 [1997]. A revised circumscription of the genera of the 
fern family Vittariaceae. Systematic Botany 22: 509-517. 


Holttum, E.H. 1955. A Revised Flora of Malaya, vol. 2., Ferns of Malaya. 
Government Printing Office, Singapore. 


Smith, A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider and P.G. 
Wolf. 2006. A classification for extant ferns. Taxon 55: 705-731. 


Smith, A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider and 
P.G. Wolf. 2008. Fern Classification, pp. 417-467. In: T.A. Ranker and 
C.H.Haufler (eds). Biology and Evolution of Ferns and Lycophytes, 
Cambridge University Press, Cambridge. 


Zhang, X-C. 2003. New combinations in Haplopteris (Pteridophyta: Vitta- 
riaceae). Annales Botanici Fennici 40: 459-461. 


Gardens’ Bulletin Singapore 62 (1): 121-126. 2010 (Dil 


A New Species of Alocasia (Araceae-Colocasieae) 
From Cambodia 


V.D.NGUYEN ,J.C. REGALADO, JR.’ AND VU TIEN CHINH! 


‘Institute of Ecology & Biological Resources, Vietnam Academy of 
Science and Technology, 18 Hoang Quoc Viet Road, Hanoi, Vietnam 
d Email: nguyenvandu@fpt.vn 
- Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, 
USA 


Abstract 


Alocasia jiewhoei V.D.Nguyen, a new species from Cambodia, is described 
and illustrated. 


Introduction 


The genus Alocasia (Araceae-Colocasieae) includes about 75 species 
distributed mainly in tropical Asia, Malesia, Australia and Melanesia 
(Govaerts and Frodin, 2002). In Indo-China, eleven species have been 
recorded in the Flore Général de |’ Indo-Chine by Gagnepain (1942). In 2007, 
during fieldwork in Cambodia, V.D. Nguyen’s group found a new species of 
Alocasia growing on dry leaf litter in forest in Kulen National Park of Siem 
Riep Province. The plant does not match any known species in the genus 
(Boyce, 2007, 2008; Hay, 1998, 1999; Hay and Wise, 1991; Medecilo et al., 
2007; Wang et al., 2005). 

A comparison of morphological characters with other species in 
the genus showed the species resemble A. odora (Lindl.) K.Koch and A. 
navicularis (K.Koch & C.D.Bouché) K.Koch & C.D.Bouché, in having 
peltate, cordate-ovate leaf blades, 5-10 pairs of lateral veins, the greenish 
spathe and conical appendices. However, it is distinguished from A. odora 
and A. navicularis by its smaller size, slender petioles, stolons with bulbils 
at the apex, conspicuous styles and prominently lobed stigmas (Table 1). It 
closely resembles A. navicularis in having erect, cymbiform spathes. 


Alocasia jiewhoei V.D. Nguyen, sp. nov. 

Alocasiae odorae similis in folii lamina peltata et spathae lamina dilute 
viridi, sed spathae lamina cymbiformi, stylis longis, stigmatibus longe lobatis 
differt.-Typus: Cambodia, Siem Riep Province, Phrom Ku Len National 
Park, 13 36’58.1” N 104 02°20.4” E, 150 m alt., V.D. Nguyen & Rattana CB- 
VN 212 (holotype, HN; isotypes, K, P, MO). Fig. 1 & Plate 1. 


122 


Table 1. A comparison of Alocasia jiewhoei and closely related species. 


Characters 


] 


A. odora 


Plant height 


40 to >250 cm 


Rhizome internal 


Light green and white 


colour 
Petiole Stout 
Bulbils on the 
None 
stolons 
Style Very short or 
i inconspicuous 
Spathe limb 
- shape Hooked cymbiform 
- colour Green to dull green 
Stigma Shallowly lobed 
Habitat Wet, evergreen forest 


Gard. Bull. Singapore 62 (1) 2010 


A. navicularis A. jiewhoei 
50 to150 cm 30 to 70 cm 
fen Ce aud Light yellow 
white 
None At the apex 


Very short or 
inconspicuous 


Erect cymbiform 


Dull yellow 


Conspicuous, 1-1.5 
mm long 


Erect cymbiform 


Medium green 


Shallowly lobed 


Wet, evergreen 
forest 


Prominently lobed 


Dry, deciduous 
forest 


Rhizomatous plants, ca 70 cm tall. Rhizomes decumbent, slender to stout, 
aerial parts 10-15 cm long, 1-4 cm in diameter, internodes 1.5-3 mm long, 
grey to dark brown externally, light yellow internally, covered by traces of 
decayed cataphylls. Roots stout, 10-15 cm long, not branched. Stolons several, 
subterranean, apex aerial, bearing bulbils at the apex; bulbils obovate, 
winged, six-angular in cross section, 1.5 cm long, ca 1 cm in diameter, oblong 
at base, abruptly acuminate, covered by several brown, small triangular 
scales. Leaves peltate, 4-7 together; petioles slender, 40-70 cm long, sheathed 
up to >1/3 of the length, green with dull dark green veins sunk below the 
surface; leaf blades ovate, 22-50 cm long, 17-30 cm at widest point, basal 
lobes broadly triangular, 7-16 cm long, 6-13 cm wide at base, apices obtuse, 
connection between 2 lobes 5-8 cm wide from petiole apex, dark green 


Alocasia jiewhoei from Cambodia 123 


Wd Z 


Figure 1. Alocasia jiewhoei V.D. Nguyen. 1. Habit; 2. Inflorescence; 3. Detail of female flower 
zone; 4. Male flower; 5. Ovary in longitudinal section; 6. Ovule. 


and glossy above, dull green beneath; lateral veins 7-8 pairs in anterior 
lobes, 2-3 veins in posterior lobes, pronounced above, prominent beneath, 
secondary veins originating from lateral veins and midrib, run parallel and 
ascend towards leaf margins, connected by reticular veins. Inflorescences 
one per axil, several per plant; peduncles slender, 25-30(-45) cm long, 7-10 
mm in diameter, expanded at top, green, covered by a thin white chalky 
layer; spathes 11-12 cm long, constricted about 1/5 of the way from the base; 
lower part ellipsoid or subglobose, green; limb 8.3-10 cm long, ca 4 cm wide, 
cymbiform, green when young, medium green at anthesis; spadices 8.5-10 cm 
long; female zones cylindrical, 0.8-1 cm long, female flowers congested, light 
yellow; ovaries subglobose in general, shallowly concaved into three lobes, 
2 mm in diameter, green; styles 1 mm long, white; stigmas three lobed, 1-1.2 
mm wide, white; ovules ca 10, placentation basal; sterile interstice conical 
at lower part, constricted near apex then slightly expanded at apex, 1.8-2.3 
cm long, 9 mm in diameter at base, 3-4 mm at the constriction and 4-5 mm 
at apex, bearing 2 whorls of white, acute-ovate, flattened sterile flowers in 


124 Gard. Bull. Singapore 62 (1) 2010 
P 


Plate 1. Alocasia jiewhoei V.D.Nguyen. A. Plant habit; B. Appendix and male portion; C. 
Staminodes and female portion in detail. (Photos: V.D. Nguyen). 


Alocasia jiewhoei from Cambodia LS) 


upper part; base covered by 2 rows of sterile flowers, columnar or ring-like; 
male zones cylindrical, 1.8-2.3 cm long, 8-10 mm in diameter, oblong to base, 
white; appendices conical, 3.5-4.8 cm long, smaller in diameter than male 
zone, acute at apex, very shallowly concaved on surface, without veins, dull 
white; male flowers sub-rectangular, 3 mm long, ca 2 mm wide, bearing 4-6 
stamens. Berries subglobose, 3-4 mm in diameter; seeds 2-3. 


Ecology: Dry seasonal forest, altitude 0-500 m. 
Distribution: Endemic in Cambodia. 
Vernacular name: None recorded. 


Uses: None recorded. However the new species has potential as an 
ornamental plant. 


Etymology: The plant is named in honour of Tan Jiew Hoe of Singapore. 


Conservation status: The plant is found in the Phrom Ku Len National Park 
of Siem Riep Province where the forest is protected. Although the new 
species is rare, there is little data to show if it is threatened. 


Acknowledgements 


In our study of the family Araceae in Indochina, the first author received 
support from the U.S. National Geographic Society Grant No. 8283-07 for 
conducting fieldwork in Laos and Cambodia and additional support from 
Mr. Tan Jiew Hoe in Singapore for the collecting expedition to Cambodia 
in 2007. The authors also thank Dr. J.F Veldkamp for the preparation of the 
Latin diagnosis and Dr. Ruth Kiew and the editor of the Gardens’ Bulletin 
Singapore for help with the English text and the constructive comments. 


References 


Boyce, P.C. 2007. Studies on the Alocasia Schott (Araceae-Colocasieae) of 
Borneo: I. Two new species from Sarawak, Malaysian Borneo. Gardens’ 
Bulletin Singapore 58: 141-154. 


Boyce, P.C. 2008. A review of Alocasia (Araceae: Colocasieae) for Thailand 
including a novel species and new species records from Southwest 
Thailand. Thai Forestry Bulletin (Botany) 36: 1-17. 


126 Gard. Bull. Singapore 62 (1) 2010 


Engler,A.andK.Krause.1920.“Additamentum adAraceas-Philodendroideas, 
Araceae-Colocasioideae”, Das Pflanzenreich T1(1V.23E): 3-132. 


Gagnepain F. 1942. Aracée. In: Lecomte, H. Flore Général de Indo-Chine 
6(9): 1075-1196. 


Govaerts, R. and D.G. Frodin. 2002. World Checklist and Bibliography of 
Araceae (and Acoraceae). Kew Royal Botanic Gardens, 560pp. 


Hay A. 1998. The genus Alocasia (Araceae-Colocasieae) in West Malesia 
and Sulawesi. Gardens’ Bulletin Singapore 50: 221-334. 


Hay A. 1999. The genus Alocasia (Araceae-Colocasieae) in the Philippines. 
Gardens’ Bulletin Singapore 51: 1-41. 


Hay, A. and R. Wise. 1991. The genus Alocasia (Araceae) in Australasia. 
Blumea 35(2): 499-545. 


Medecilo MP, G.C. Yao and D.A. Madulid. 2007. A new species of Alocasia 
(Araceae: Colocasieae) from Panay Island, Philippines. Journal of the 
Botanical Research Institute of Texas 1(2): 815-818. 


Wang, Yue-Hua, Jian-Tao Yin and Zai-Fu Xu, 2005. Alocasia hypnosa 
(Araceae), a new species from Yunnan, China. Annales Botanici Fennici 
42: 395-398. 


Gardens’ Bulletin Singapore 62 (1): 127-133. 2010 127 


New Combinations in Malaysian Staphyleaceae 


A.T. NOR-EZZAWANIS 


Forest Research Institute of Malaysia 
52109 Kepong, Selangor, Malaysia 


Abstract 


New combinations are made for the seven species of Dalrympelea 
(Staphyleaceae) from Malaysia that were previously included in Turpinia. 
Dalrympelea pomifera Roxb. is distinct from D. sphaerocarpa (Hassk.) 
A.T.Nor-Ezzawanis and does not occur in Malaysia and Turpinia ovalifolia 
‘Elmer from the Philippines is distinct from D. trifoliata (Ridl.) A.T.Nor- 
Ezzawanis. 


Introduction 


Staphyleaceae currently includes two genera, Dalrympelea Roxb. and 
Staphylea L. (Simmons, 2007; Stevens, 2010), that can be distinguished by 
the following morphological characters: stoloniferous shrubs (2 m) to upper 
canopy trees of 25-30 m in Dalrympelea, and small trees reaching 15 m in 
Staphylea; the bark ranges from creamy yellow and flaky to smooth grey 
in Dalrympelea, and grey to black and somewhat mottled, with or without 
lenticels in Staphylea: the stipules of opposed leaves are fused and sometimes 
becoming bifid at the apex, often having a colleter in Dalrympelea, and the 
stipules are free and multi-veined in most species of Staphylea. The former 
occurs in the Old World and the latter in both the Old and New Worlds. 
Based on molecular analysis using nuclear and chloroplast markers, 
Staphyleaceae divides into two clades that are treated as distinct at the 
generic level (Simmons & Panero, 2000; Simmons, 2007). One clade, 
Staphylea, includes all species of Staphylea, all New World species of the 
Turpinia Raf.,and one Asian species, Turpinia cochinchinensis (Lour.) Merr. 
and the monotypic Euscaphis japonica (Thunb.) Kanitz (Simmons. 2007). 
The other clade includes the Old World Turpinia species, except 
for Turpinia cochinchinensis. Members of this latter clade are generally 
characterized by being evergreen trees with fused stipules (at least at the 
base), coriaceous leaves and a berry with a thickened pericarp (Simmons. 
2007). Because the type species of Turpinia is T. cochinchinensis, which is 
now included in Staphylea,the name Turpinia cannot be applied to the Asian 
species. The earliest name for this clade is Dalrympelea Roxb. (Roxburgh, 


128 Gard. Bull. Singapore 62 (1) 2010 


1819). The revision of the family for the Flora of Peninsular Malaysia 
necessitates making new combinations for local Turpinia species besides 
reassessing the status of some species. New combinations are also provided 
for species in the Malaysian states of Sabah and Sarawak in Borneo that 
were revised by Pereira (1995). 


DALRYMPELEA Roxb. 

Hort. Beng. (1814) 17, nomen, Pl. Corom. 3 (1819) 76, t. 279; Simmons in 
Kubitzki, Fam. Gen. Vasc. Pl. 9 (2007) 443. Type species: Dalrympelea 
pomifera Roxb., Pl. Corom. 3 (1819) 76. ¢. 279. 

Synonym: Turpinia Vent., Choix (1803) 31, t.31, p.p. excl. New World species 
and T. cochinchinensis; Ridley, Fl. Malay Pen. 1 (1922) 511; Linden, FI. 
Malesiana 1, 6 (1960) 51; Whitmore, Tr. Fl. Malaya 1 (1972) 447; Pereira, Tr. 
Fl. Sabah & Sarawak (1 (1995) 454. 


Evergreen shrubs to large trees to 30 m tall, sometimes with buttresses. 
Leaves pinnately (1- or 3-) or 5-15-foliolate, coriaceous; stipules interpetiolar, 
basally connate to fused along their length. Inflorescences terminal or 
axillary panicles with more than 100 flowers. Flowers with sepals free, shorter 
than the pink, cream, yellow or green-white petals; stamens arising between 
the lobes of the nectary disc; ovary (2-)3(-4)-locular, sometimes partially 
imbedded in the disc, more or less syncarpous, with 2-8 ovules per locule. 
Fruit an ellipsoid, nearly globose or trilobed berry, fleshy or leathery, usually 
green to purple; exocarp thick and fleshy to woody. Seeds 1-6 per fruit. 


Distribution: About 20-25 species from Sri Lanka and S India to China and 
Japan and southwards to New Guinea (Simmons, 2007) with 8 species in 
Malaysia (2 species in Peninsular Malaysia and 6 species and | variety in 
Sabah and Sarawak). 


New Combinations for Malaysian Species 


1. Dalrympelea borneensis (Merr. & L.M.Perry) A.T.Nor-Ezzawanis, comb. 
nov. 

Basionym: Turpinia montana var. borneensis Merr. & L.M.Perry, J. Arnold 
Arbor. 22 (1941) 553. -Homotypic synonym: Turpinia borneensis (Merr. & 
L.M.Perry) B.L.Linden, Fl. Malesiana 1, 6 (1960) 56; Pereira, Tr. Fl. Sabah 
& Sarawak 1 (1995) 456. -Type: Borneo, Sabah [British North Borneo], 
Gunung Kinabalu, Tenompok, J. & M.S. Clemens 29391 (holotype, UC; 
isotypes, BO, K). 


Distribution: Borneo (Kalimantan and Sabah) and the Philippines. 


New Combinations in Malaysian Staphyleaceae 129 


2. Dalrympelea calciphila (J.T.Pereira) A.T.Nor-Ezzawanis, comb. nov. 
Basionym: Jurpinia calciphila J.T.Pereira, Sandakania 5 (1994) 18, fig. 1, Tr. 
Fl. Sabah & Sarawak 1 (1995) 457. -Type: Borneo, Sarawak, Miri Division, 
Gunung Api, Anderson S 4710 (holotype, SAR; isotypes, BO, K, L, SING). 


Distribution: Endemic in Sarawak, Borneo — Kuching Division near Bau 
(Bukit Buan and Bukit Gebung) and Miri Division (Gunung Api and 
Gunung Buda). 


3. Dalrympelea grandis (B.L.Linden) A.T.Nor-Ezzawanis, comb. nov. 
Basionym: Turpinia grandis B.L.Linden, Fl. Malesiana 1,6 (1960) 55; Pereira, 
Tr. Fl. Sabah & Sarawak 1 (1995) 457. -Type: E Borneo, W Koetai, Kiau 
River, Endert 4669 (holotype, L; isotypes, BO, K). 

Distribution: Endemic in Borneo (Kalimantan, Sabah and Sarawak). 


4. Dalrympelea nitida (Merr. & L.M.Perry) A.T.Nor-Ezzawanis, comb. nov. 
Basionym: TJurpinia nitida Merr. & L.M.Perry, J. Arnold Arbor. 22 (1941) 549; 
Linden, Fl. Malesiana 1, 6 (1960) 58; Pereira, Tr. Fl. Sabah & Sarawak 1 
(1995) 459.-Type: Borneo, Sabah [British North Borneo], Gunung Kinabalu, 
Penibukan, J. & M.S. Clemens 30840 (holotype, UC; isotypes, BO, K). 


Distribution: Endemic in Sabah, Borneo (Ranau, Sandakan, Kudat and 
Lahad Datu Districts). 


5. Dalrympelea sphaerocarpa (Hassk.) A.T.Nor-Ezzawanis, comb. nov. 
Basionym: Turpinia sphaerocarpa Hassk., Flora 25, 2 (1842) Beibl. 1, 42: 
Ridley, J. Str. Br. Roy. As. Soc. 82 (1920) 179; Linden, Fl. Malesiana 1, 6 
(1960) 49; Whitmore, Tr. Fl. Malaya 1 (1972) 448; Pereira, Tr. Fl. Sabah & 
Sarawak 1 (1995) 460-Homotypic synonym: Turpinia pomifera (Roxb.) DC. 
var. sphaerocarpa (Hassk.) King, J. As. Soc. Beng. 65 (1896) 453. —Type: Java. 
sine coll., s.n., Herb. Reinwardtianum (holotype, L, acc. no. 908272875). 
Heterotypic synonyms: Turpinia latifolia Wall. ex Ridl., J. Str. Br. Roy. As. 
Soc. 82 (1920) 178, Fl. Mal. Pen. 1 (1922) 512Type: Singapore, Wallich 4939 
(lectotype, K, isotype, CAL). -Turpinia laxiflora Ridl., J. Str. Br. Roy. As. Soc. 
82 (1920) 179, FI. Malay Pen. 1 (1922) 512; Linden, Fl. Malesiana 1, 6 (1960) 
57.—-Type: Peninsular Malaysia, Perak, Larut, Kunstler 2824 (holotype, K). 


Taxonomic notes: Wallich (1828) in his Catalogue named Wallich 4939 as 
‘Turpinia ?latifolia’. Hiern (1875) included Turpinia latifolia as a synonym 
of Turpinia pomifera (Roxb.) DC. with Wallich 4939 as the only specimen 
cited for Malaya. 


130 Gard. Bull. Singapore 62 (1) 2010 


King (1896) described two varieties of Turpinia pomifera from 
Peninsular Malaysia. The typical variety was represented by asingle specimen, 
Kings’ Collector 4243 from Perak, and Turpinia pomifera var. sphaerocarpa 
(Hassk.) King was described as ‘the common form in the Malayan Provinces’. 
King cited 7: latifolia based on Wallich 4939 as asynonym of var. sphaerocarpa. 

Turpinia sphaerocarpa Hassk. was originally described from Java 
and has small fruits 0.5-1.8 cm diameter with a thin pericarp 1-4 mm thick 
as compared with Turpinia pomifera from Continental Asia that Linden 
(1960) described as having larger fruits 2.5-3.7 cm diameter with a pericarp 
5 mm thick. Re-constructing the fragments of the fruit on Kings’ Collector 
4243 shows that it has a small fruit about 1.5 cm diameter with a pericarp 
ca 3 mm thick showing conclusively that it is not a specimen of Turpinia 
pomifera, but is typical of Turpinia sphaerocarpa. Ridley (1920) was not 
able to examine Kings’ Collector 4243 (there is apparently no specimen at 
Kew) but considered it was unlikely to be the same as the Turpinia pomifera 
and instead he used Wallich’s original name, T- latifolia Wall. ex Ridl., to 
describe this common species in Peninsular Malaysia. (Incidentally, he gave 
Penang Hill as the locality for Wallich 4939 but in fact Wallich collected it 
from Singapore). Ridley’s description mentioned the fruit diameter as ca 
1.3 cm (within the Turpinia sphaerocarpa range), but unfortunately he did 
not record pericarp thickness. Linden (1960) too was doubtful that Turpinia 
pomifera occurred in Peninsular Malaysia and considered Kings’ Collector 
4243 as "too inadequate" for identification. He therefore reinstated Turpinia 
sphaerocarpa to specific rank with Turpinia pomifera var. sphaerocarpa and 
T. latifolia as synonyms. Whitmore (1972) noted that Turpinia pomifera was 
based‘onone doubtfulcollection’, namely Kings’ Collector 4243. Itis therefore 
now clear that Turpinia pomifera does not occur in Peninsular Malaysia. 

Of the three type specimens at L (acc. nos. 908272875, 908272863 and 
908272865), sheet number 908272875 is chosen as the lectotype because it 
is the most complete specimen with more fruits and it is from Herbarium 
Reinwardtianum. 

Ridley (1920) described a new species of Turpinia, T laxiflora Ridl., but 
without a description of the fruit. In his Flora (Ridley, 1922), he distinguished 
it from 7: latifolia by panicle size: 20 cm [8 in] long in T: latifolia as opposed 
to 38 cm [15 in] in 7: laxiflora. Linden (1960) followed Ridley in retaining 
T: laxiflora as a distinct species noting that it differed from 7. sphaerocarpa 
in its wider leaflets (8-24 x 3.5-12 cm as opposed to 3-18 x 2-10 cm in 7. 
sphaerocarpa), larger panicles (to 45 cm vs. to 30 cm long), narrower petals 
(2.25-2.5 x 1-1.25 mm vs. 2.5 x 1.5 mm), shorter filaments (1.5-1.75 mm vs. 
2-2.5 mm long), smaller fruits (0.5-1 cm vs. 1-1.5 cm diameter) and very thin 
pericarp (0.2-0.9 mm thick vs. 1-3 mm thick in 7: sphaerocarpa). However, 
Whitmore (1972) synonymised T: laxiflora with T. sphaerocarpa but without 
giving a reason for his decision. 


New Combinations in Malaysian Staphyleaceae 13] 


Examination of specimens from Peninsular Malaysia identified as 7. 
latifolia/T. sphaerocarpa and T. laxiflora show that the characters listed by 
Linden (1960) are not mutually exclusive. For example, Corner 28690 (KEP) 
has a panicle ca 21 cm long, which falls within Dalrympelea sphaerocarpa, 
but the petals measure ca 2 x 1.25 mm and filaments ca 1.25-1.5 mm long 
and so fall within the range for T: laxiflora. Similarly, Jaamat 10275 (KEP) 
has a fruit ca 1.2 cm diameter with a pericarp ca 4 mm thick that fits D. 
sphaerocarpa but the petals measure ca 2 x 1 mm and the filaments are ca 
1.5 mm long that fall within the range of T: laxiflora. Linden (1960) also 
mentioned colour of the dried leaf as a distinguishing characters (brown on 
both surfaces in T: laxiflora, but discolorous in D. sphaerocarpa), but again it 
was found that this character did not distinguish these two taxa. There being 
no constant character or combination of characters to keep 7: laxiflora apart, 
it is here treated as a synonym of D. sphaerocarpa. 

Burkill (1966) confused the picture by recognizing Turpinia pomifera 
and treating both 7. Jatifolia and T. laxiflora as synonyms of it. He described 
T. pomifera as a “fairly large tree in the lowlands” which indicates that he 
was probably referring to Dalrympelea sphaerocarpa. 

Pereira (1994) recognized a second variety, var. microcerotis, from 
Borneo, which differs from var. sphaerocarpa in having shorter leaf rachises 
(2-8 cm vs. 10-18 cm long), shorter leaflets (5-10 cm vs. 7-19 cm long) and 
a slightly trilobed fruit with three apical horns compared with the globose, 
hornless fruit of var. sphaerocarpa. 


Distribution: Sumatra, Peninsular Malaysia, Singapore, Borneo, Java, Lesser 
Sunda Islands (Bali, Flores), Sulawesi, the Philippines and Maluku (Ceram, 
Ambon). 


5a. Dalrympelea sphaerocarpa (Hassk.) A.T. Nor-Ezzawanis var. 
microcerotis (J.T. Pereira) A.T.Nor-Ezzawanis, comb. nov. 

Basionym: Turpinia sphaerocarpa Hassk. var. microcerotis J.T. Pereira, 
Sandakania 5 (1994) 21, fig. 2; Tr. Fl. Sabah & Sarawak 1 (1995) 461. — 
Type: Borneo, Sabah [British North Borneo], Ranau, Gunung Kinabalu, 
Tenompok, J. & M.S. Clemens 28707 (holotype, SING; isotypes, BO, K). 


Distribution: Endemic in Borneo (Sabah and Kalimantan). 


6. Dalrympelea stipulacea (B.L.Linden) A.T.Nor-Ezzawanis, comb. nov. 
Basionym: Turpinia stipulacea B.L.Linden, Fl. Malesiana 1, 6 (1960) 55; 
Pereira, Tr. Fl. Sabah & Sarawak 1 (1995) 461.—Type: Borneo, Sabah, Gunung 
Kinabalu, near Tibabah River, Carr SFN 27516 (holotype, SING). 
Distribution: Endemic to Sabah (Ranau district only). 


132 Gard. Bull. Singapore 62 (1) 2010 


7. Dalrympelea trifoliata (Ridl.) A.T.Nor-Ezzawanis, comb. nov. 
Basionym: Turpinia trifoliata Ridl., J. Str. Br. Roy. As. Soc. 82 (1920) 178, FI. 
Malay Pen. 1 (1922) 511, fig. 50. -Type: Peninsular Malaysia, Melaka, Nyalas, 
Goodenough 1771 (lectotype, SING, here chosen). 

Heterotypic synonym: Turpinia ovalifolia Elmer, Leafl. Philip. Bot. 2 (1908) 
490, auct. non: Linden, Fl. Malesiana 1,6 (1960) 58; Whitmore, Tr. Fl. Malaya 1 
(1972) 448.-Type: Philippines, Luzon, Lucban Tayabas, Elmer 8088 (SING). 


Taxonomic notes: Some authors (Linden, 1960; Whitmore, 1972) considered 
this species in Peninsular Malaysia to be conspecific with Turpinia ovalifolia. 
However, careful comparison between Peninsular Malaysian specimens 
and the type specimens of both 7. ovalifolia and T. trifoliata show that the 
Malaysian specimens previously identified as 7. ovalifolia in fact belong 
to T. trifoliata. Turpinia ovalifolia, known only from the type specimen, is 
characterized by its small leaves ca 4 x 2.5 cm with a more-or-less entire 
margin. In contrast, T° trifoliata specimens cited by Ridley (1920), namely 
Goodenough 1771 and Ridley 15906, are characterized by larger leaves ca 7- 
14 x 3.5-7 cm with a crenate margin with fine, dark brown (when dry) callous 
points. These characters are shared by more recently collected specimens 
from Malaysia and indeed some have even larger leaves up to ca 9.5-20 
x 4-9.5 cm. Therefore it is concluded that 7: ovalifolia does not occur in 
Peninsular Malaysia. 

Ridley (1920) cited two specimens in his original description of the 
species. Goodenough 1771 is here chosen as the lectotype because it is a 
more complete specimen and the sheet has Ridley’s description of the 
flower (in his handwriting) and is named ‘7urpinia trifoliata’. 


Distribution: Peninsular Malaysia (Perak, Selangor, Negeri Sembilan, 
Pahang and Johor). 


Acknowledgements 


This study was carried out for the Flora of Peninsular Malaysia Project 
funded by the Ministry of Science, Technology and Innovation (MOSTI) 
through the National Council for Scientific Research and Development 
(MPKSN) under Project No. 01-04-01-0000 Khas 2 entitled ‘Safeguarding 
the Forest Plant Diversity of Peninsular Malaysia’. I am indebted to P.F. 
Stevens for advice on the current status of Staphylea and Dalrympelea and 
to Drs. R. Kiew, Saw L.G., R.C.K. Chung, and E. Soepadmo for help with 
preparing the manuscript. I should also like to thank the Curators of the 
herbaria at KLU, UKMB and SING for loan of specimens and to BM, K and 
L for access to their collections. 


New Combinations in Malaysian Staphyleaceae 133 


References 


Burkill, 1.H., 1966. Turpinia pomifera. In: A Dictionary of the Economic 
Products of the Malay Peninsula. p. 2234. 


Hiern, W.P. 1875. Turpinia. In Hooker,f Flora of British India 1: 698-699. 
King, G. 1896. Turpinia. Journal of the Asiatic Society of Bengal 62: 452-453. 
Linden, B.L. van der. 1960. Staphyleaceae. In: Flora Malesiana 1 (6): 49-59. 


Pereira,J.T.1994. Anewspecies andanewvariety of Turpinia (Staphyleaceae) 
from Borneo. Sandakania 5: 15-23. 


Pereira, J.T. 1995. Staphyleaceae. In: Tree Flora of Sabah & Sarawak 1: 453- 
462. 


Ridley, H.N. 1920. New and rare species of Malayan plants. Journal of the 
Straits Branch of the Royal Asiatic Society 82: 177-179. 


Ridley, H.N. 1922. Turpinia. In: Flora of the Malay Peninsula 1: 511-512. 


Roxburgh, W. 1819. Dalrympelea pomifera. In: Plants of the East Coast of 
Coromandel, pp. 76-77, t. 279. 


Simmons, S.L. 2007. Staphyleaceae. In: Kubitzki, K.(ed.). The Families and 
Genera of Vascular Plants 9: 443-445. 


Simmons, S.L. and J.L. Panero. 2000. Phylogeny and biogeography of 
Staphyleaceae (DC.) Lindl. American Journal of Botany 87(6, suppl.): 
S77. 


Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, 
June 2008. http://www.mobot.org/MOBOT/research/A Pweb/. 
Latest update January 2010. 


Wallich, N. 1828. Turpinia latifolia. In: Numerical List of Dried Specimens of 
Plants, p. 173. 


Whitmore, T.C. 1972. Staphyleaceae. In: Tree Flora of Malaya 1: 446-448. 


dir? @& que 7 
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2 ( ae 
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Gardens Bulletin Singapore 62 (1): 135-142. 2010. 135 


A New Species of Costaceae from Borneo 


A.D.POULSEN’ AND C.D. SPECHT” 


Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh 
: EH3 5LR, Scotland, U.K. 
- Plant and Microbial Biology, University of California, Berkeley, 
111 Koshland Hall, MC 3102, Berkeley, CA 94720, USA 


Abstract 


A new species, Cheilocostus borneensis, is described. Specimens were 
collected in Sarawak in 1987 and Kalimantan in 2000, but only intensified 
surveys of gingers in Sarawak in 2002-2004 provided sufficient collections 
to recognize the new species, which is here described and illustrated. It is 
closely related to the widespread C. globosus from which it differs by the 
chocolate-brown sheaths, absence of axillary shoots on vegetative stems, 
larger leathery leaves, and by its calyx that is not prickly. 


Introduction 


Members of Bornean Costaceae were previously placed in the genus 
Costus L. (Maas, 1979). which is now circumscribed as a clade comprising 
only African and neotropical species based on phylogenetic analyses of 
morphological and molecular data (Specht and Stevenson, 2006). Following 
this evaluation of the generic circumscription of Costaceae, only two genera, 
Cheilocostus C. Specht and Paracostus C. Specht, are native to Borneo where 
seven species of Costaceae are presently known (Maas. 1979: Meekiong 
et al., 2006; Meekiong et al., 2008). The exact generic placement has not 
been established for all Bornean species and an updated revision for both 
Cheilocostus and Paracostus is pending and will likely include other recently 
discovered and described species. 

The genus Cheilocostus is easily distinguished from Paracostus 
by consisting of larger plants (> 1.5 m high) with erect shoots, and a 
condensed inflorescence with conspicuous bracts, each subtending a single 
flower. Paracostus, in contrast, is characterized by smaller plants (< 1.5 
m), prostrate stems with few leaves, and inflorescences with few flowers 
supported by inconspicuous bracts. Cheilocostus is closely related to the 
genus Japeinochilus Mig. which is only found east of Sulawesi into New 
Guinea, Australia and the Pacific (Gideon, 1996; Poulsen er al. 2010). 


136 Gard. Bull. Singapore 62 (1) 2010 


During expeditions targeting gingers in Sarawak in 2002-2004, the 
first author collected, with Malaysian collaborators, new material of several 
species of Costaceae. One of these had already been collected several times, 
and specimens were deposited in several herbaria but without pickled 
flowers essential for its description. The material now being available, the 
species is described below. 


Cheilocostus borneensis A.D. Poulsen, sp. nov. 

Cheilocostus borneensis in inflorescentia radicaliC. globoso similis est 
sed ab eo foliis ad apicem caulis aggregatis plerumque majoribus et calyce 
molliter acuto (haud pungenti nec aculeato) differt. -Typus: Malaysia, Borneo, 
Sarawak, Batang Ai, Sungai Senkabang, small stream connecting to Sg. 
Delok opposite of Ng. Sumpa longhouse, 1°12’S 112°3’E, 130 m, flowering 
8 Dec 2002, A.D. Poulsen & Bakir Raymond 1964 (holo, SAR; iso, AAU, 
Sarawak Biodiversity Centre Flora Depository). Figs. 1 & 2. 


Figure 1. Cheilocostus borneensis photographed by A.D. Poulsen (Poulsen 2696, cultivated). 


A New Species of Costaceae from Borneo 137 


Terrestrial, perennial herb. Leafy shoots in a dense clump, 1.5-2 m tall. Base of 
leafy shoot to 3 cm diam., pale yellow-green when fresh, covered by reddish 
brown sheaths. Stem leafless in lower ca 1.4 m, sheaths reddish chocolate- 
brown (uppermost yellow-green), glabrous, with 6-10 leaves consistently 
clustered toward the shoot apex. Ligule 1-2 mm long, + truncate, slightly 
longer laterally to the petiole. Petiole 5-12 mm long, swollen, pale yellowish, 
slightly canaliculate, glabrous. Lamina to 27-42 x 9-16(-20) cm, narrowly 
obovate, mid-green above, pale green beneath, coriaceous, slightly plicate, 
glabrous throughout, base narrowly cuneate, apex acuminate, ca 1 cm long. 
Inflorescence radical (i.e. at the base of the plant terminating a separate 
leafless shoot that emerges directly from the rhizome), 12-21 cm long (bracts 
only), lax. Peduncle horizontal to ascending, 6-12 cm long, sheaths + tubular, 
brown, glabrous, margin ragged. Spike 4-12 x 5-8 cm (bracts only). Fertile 
bracts to 3.2-3.5 x 2-2.5 cm, elliptic, cucullate, margin membranous, apex 
+ emarginate, softly mucronate, dark brown to greenish or pale reddish 
brown, glabrous. Rachis (distance from base of lowermost ovary to base 
of uppermost ovary) 2 cm long extending with age to 7(-9) cm, with 40-80 
flowers, 1-2 open at a time. Bracteole 2-2.5 cm long, split to base adaxially, 
reddish brown, glabrous, apex rounded with one minute mucro, 1-lobed 
developed laterally, cucullate, not closely adhering to calyx, sometimes with 
a second shorter lobe without mucro. Flower 8.5-10 cm long, exerted ca 5 
cm above the supporting bract. Calyx 3-3.2 cm long, tubular, dark reddish 
brown, glabrous, apex 3-lobed, lobes 0.7-1 cm long, slightly involute, apex 
acute to mucronate, soft (not pungent). Corolla tube (from apex of ovary 
to base of divergence of labellum and stamen) 2.5-3.2 cm long, fused solid 
with style in lowest 1-1.5 cm, white at base; lobes 3.5 x 1.5 cm, narrowly 
obovate, translucent white, glabrous, apex rounded, finely apiculate to 1 
mm long. Labellum tube (from insertion of dorsal corolla tube to base of 
divergence of labellum and stamen) 1.1-1.3 cm long with coarse yellow hairs 
inside and outside. Labellum 5-5.3 x 5.2-6 cm, broadly obovate, thickened 
in centre with course, short hairs, white with yellow center, margin finely 
undulating, glabrous. Stamen ca 2.5 cm long (ca 3.2 cm when crest flattened), 
ca 1 cm wide, petaloid, white. Anther crest slightly or irregularly 3-lobed, 
recurved, 1.1-1.2 cm long, with yellow spot in lower centre, coarse hairs at 
margin. Thecae 0.7-0.8 cm long, 0.3-0.4 cm across both, dehiscing for their 
entire length. Ovary 1 x 0.6 cm, flattened ellipsoid, glabrous. Style 3.5-3.7 
cm long (free part), glabrous. Stigma ca 0.3 cm wide, fan-shaped, flattened 
2-lobed, lobes overlapping, one larger than the other, hairy, pale yellow. 
Infructescence head to 12 x 8 cm, often still flowering at apex, with persistent 
bracts, bracteoles and, calyces. Fruit 0.9-1 x 0.8-0.9 cm, obovoid, flattened- 
triangular, with an apical column (base of calyx and corolla tube), 3-locular, 
glabrous, cream to pale green. Seeds 1.5-1.7 x 1-1.2 mm, irregularly barrel- 
shaped, black, aril white, basal and not enclosing the seed. 


138 Gard. Bull. Singapore 62 (1) 2010 


£ 


& 


fb fl br ca 


Plate 1. Cheilocostus borneensis. A. Habit; B. Leaves; C. Infructescence with flowers at apex; 
D. Close up of labellum and stamen; E. fb = fertile bract, fl = fertile bract with single flower, br 
= bracteole, ca = calyx, co = flower with bracteole and calyx removed, tr = longitudinal section 
of flower, sd = stamen, dorsal view, sv = stamen, ventral view; F. Fruit with semipersistent 
calyx. Photographs by A.D. Poulsen, A, C—D, F of Poulsen & Bakir Raymond 1964 (the type): 
B, E of Poulsen 2696 (cultivated). 


A New Species of Costaceae from Borneo 139 


Additional materials examined: INDONESIA. Borneo. West Kalimantan. 
Camp Betung Kerihun NP, Putussibau, 0°56’N 113°19E, 150 m, fruiting 28 
Feb 2000, Ambriansyah, Kade Sidiyasa & Albertus AA 2238 (BO, L, WAN). 
MALAYSIA. Borneo, Sarawak: Batang Ai, Sungai Senkabang, small stream 
connecting to Sg. Delok opposite of Ng. Sumpa longhouse, 1°12’S 112°3’E, 
130 m, flowering 3 Jun 1993, Christensen & Poulsen 1997 (AAU): Gunung 
Mulu National Park, R. Ubong, between Mulu N.P. and logging concession, 
near Base Camp, 200 m, flowering, 7 Nov 1990, Warwick MW177 (E bar code 
E00128356): Kapit, Balleh, Ulu Sungai Mengiong, Wong Kijang, flowering 26 
Oct 1988, Othman et al. S.56077 (SAR): Kapit, Balleh, Ulu Sungai Mengiong, 
Nanga Sebaning, 1°25’N 113°25’E (indicated approximately on map on 
rear side of label), flowering 1 Nov 1988, Othman et al. 8.56464 (AAU, E, 
K n.v.); Kapit, Batang Baleh, Sungai Mengiong, Sungai Entulu, 18 Jul 1987. 
Bernard Lee. S. 54624 (AAU, E (bar code E00320502: bar code E00304605), 
K n.v.); Cultivated at Royal Botanic Garden Edinburgh, Accession number 
and qualifier: 20040728*A, collected 16 Aug 2007, Poulsen 2596 (E, SING), 
origin Poulsen & Bakir Raymond 1964 (the type). 


Distribution: Endemic to Borneo where it is known from three main areas 
in Sarawak (one being three collections from the Kapit area) and one in 
Kalimantan. The furthest localities are about 450 km apart. 


Local names and uses: Pa’bu (Iban language; Poulsen & Bakir Raymond 
1964). The plant was used in the past by Iban people but it is not certain for 
what purpose; pasat baju (Iban; Othman et al. 8.56077). 


Etymology: The epithet refers to the species being endemic to Borneo. 


Ecology and habitat. Lowland primary or secondary (logged) mixed 
dipterocarp forest, along riverbanks, at 130-200 m. 


Conservation status: C. borneensis is found at the foot hills of the central 
mountain range of Borneo in an area covering at least 2200 km” but with 
fewer than 10 localities. Also its sexual reproduction seems dependent on 
natural pollinators that may not persist in degraded habitats (see Notes 
below). Currently we propose the category of Vulnerable (Vu Blab(iii); 
IUCN, 2001), but because of the logging activities and land use is changing 
rapidly in Borneo, this category could soon change to “Endangered”. 


Notes: Cheilocostus borneensis deviates from the generic description (Specht 
and Stevenson, 2006) by not having axillary branching of the vegetative 


140 Gard. Bull. Singapore 62 (1) 2010 


fi02 115° 


Figure 2. Distribution of Cheilocostus borneensis presently known from Sarawak (Malaysia) and 
West Kalimantan (Indonesia). 


shoots. Vegetative branching is shared by Cheilocostus and Tapeinochilos 
but is lost in some Tapeinochilos species and, similarly, may have been lost 
in C. borneensis. Molecular evidence using four gene regions (ITS, ETS, 
rpb2 and trnL-F) and samples from a single accession (the type) places this 
species in Cheilocostus, sister to C. globosus (Blume) C. Specht. 
Cheilocostus borneensis is similar to C. globosus in having a radical 
inflorescence, but is easily distinguished by having distinctly chocolate- 
brown sheaths on the stem (also obvious in herbarium specimens), having 
few larger leaves congested at the top of the stem, the leathery almost 
plastic-like texture of the leaves, and by not having axillary branching. As 
the flowers of C. globosus vary in colour of the labellum from white to 
yellow, dark orange to red, this character by itself is not always conclusive 
in separating the two. A more useful floral character to separate the two 
species is the apex of the calyx: C. borneensis is soft, whereas C. globosus has 
rigid points that, if touched, will easily make you bleed (i.e. pungent). 
Recently, Costus mulus Meekiong, Ipor & Tawan was described from 
Sarawak (Meekiong et al., 2008). This species also has a radical inflorescence 
containing flowers with a white labellum but the yellow patch is confined to 
a dot at the terminal margin of the labellum. Even though the description 
leaves many questions open and we have not been able to examine the type 


A New Species of Costaceae from Borneo 14] 


material, this species has smaller leaves (13-21 x 6.5-9.5 cm vs. 27-42 x 9- 
15 cm) with a rounded to cordate base (not narrowly cuneate) and thus is 
clearly distinct from Cheilocostus borneensis. 

The northernmost locality presently known of C. borneensis is Mulu 
National Park in Sarawak. Despite Smith’s inventory (1984) on the richness 
of gingers of this area, no material was collected during her studies. Thus. she 
only reported Paracostus paradoxus (K.Schum) C. Specht and Cheilocostus 
speciosus (J. KOnig) C. Specht from this area. The collection from Mulu. 
Warwick MW177, is a convincing match to the other collections. but the 
mucro on the bracts are exceptionally prominent. 

In nature, the spike of Cheilocostus borneensis may have mature fruits 
in the lower bracts while simultaneously flowering towards the apex (Plate 
1C). In cultivation mature fruits were never observed to develop and it is 
likely that manual pollination is needed to produce seeds. This may indicate 
that C. borneensis is only able to set fruits in habitats where the natural 
pollinator is present. 

On 11 Dec 2002. at Sungai Rirang near the type locality, a plant of 
Cheilocostus borneensis with normal leafy shoots was seen bearing an erect, 
54 cm long peduncle that was similar in appearance to the chocolate-brown 
sheathed stem of a leafy shoot, but instead of bearing leaves it terminated 
in a spike. This aberrant inflorescence has only been seen this once, never 
occurred in cultivation, and is apparently a rare phenomenon. 


Acknowledgements 


The first author thanks his adopted family at Nanga Sumpa for help 
with the fieldwork during which this species was discovered, Sarawak 
Biodiversity Centre and the National Parks and Nature Reserves of 
Sarawak for processing permits, HRH Crown Prince Frederik’s Foundation 
for sponsorship of fieldwork, and Paul and Hiltje Maas for confirming the 
species as new during a visit to Utrecht in 2003. We thank Robert Mill and 
Philip Oswald for help with the Latin diagnosis and John Mood, as well as 
two anonymous reviewers, for critically reading an earlier version of the 
manuscript. 


References 


Gideon, O.G. 1996. Systematics and evolution of the genus Tapeinochilos 
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Doctor of Philosophy. Department of Botany and Tropical Agriculture, 
James Cook University. Queensland, Australia. 385 pp. 


142 Gard. Bull. Singapore 62 (1) 2010 


IUCN 2001. IUCN Red List Categories and Criteria. Version 3.1. IUCN 
Species Survival Commission. Gland, Switzerland and Cambridge, UK. 


Maas, PJ.M. 1979. Notes on Asiatic and Australian Costoideae 
(Zingiberaceae). Blumea 25: 543-549. 


Meekiong, K., Ipor, I.B. and C.S. Tawan. 2006. Three new species from 
Sarawak, and notes on the cauliferous Costus Group (Costaceae, 
subgenus Paracostus). Folia Malaysiana 7: 55-72. 


Meekiong, K., Ipor, I.B., Tawan, C.S. and M. Muliati. 2008. A new Costus 
(Costaceae) from Sarawak, Borneo, Malaysia. Rheedea 18: 87-89. 


Poulsen, A.D., Gideon, O.G. and M. Ardiyani. 2010. Names of Tapeinochilos 
(Costaceae) in Wallaceae. Blumea 55: 61-64. 


Smith, R.M. 1984. Zingiberaceae of Mulu National Park, pp. 85-96. In: A.C. 
Jermy (ed.), Studies on the flora of Gunung Mulu National Park, Sarawak. 
Kuching, Sarawak. 


Specht, C.D. and D.W. Stevenson. 2006. A new phylogeny-based generic 
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Gardens’ Bulletin Singapore 62 (1): 143-153. 2010 143 


Three New Species of Begonia Sect. Platycentrum 
from Nepal 


S.RAJBHANDARY .M. HUGHES’ AND K.K.SHRESTHA | 


Central Department of Botany 
Tribhuvan University. Kirtipur, Kathmandu, Nepal 
Royal Botanic Garden Edinburgh 
20A Inverleith Row, Edinburgh, U.K.EH3 5LR 
Author for correspondence: S. Rajbhandary:imogine3@gmail.com 


Abstract 


Three new species of Begonia (Begoniaceae) are described from Nepal. All 
three species (Begonia nuwakotensis S. Rajbhandary. Begonia panchtharensis 
S. Rajbhandary and Begonia taligera S. Rajbhandary) belong to Begonia 
section Platycentrum, and they increase the number of Begonia species 
known from Nepal to 22. All are considered to belong to the IUCN threat 
category VUD2. 


Introduction 


Begonia inhabits moist, shady locations in humid lowland to upland forests. 
The greatest number of species is found in mid-elevations ranging between 
1200-2500 m, growing in cloud forest habitats in rock crevices and on moist 
boulders and moss-covered tree trunks. Begonia picta Sm. and Begonia 
dioica Buch.-Ham. ex D. Don are the most common species in Nepal, found 
growing from 150-2700 m altitude. The genus was previously represented 
by 18 species in Nepal (Hara er al., 1978; Doorenbos er al., 1998: Press et 
al., 2000) and one new record (Rajbhandary and Shrestha, 2009) bringing 
this total to 19. These are placed within five different sections, Diploclinium 
(Lindl.) A.DC.. Monopteron (A.DC.) Warb., Platycentrum (Klotzsch) 
A.DC, Putzeysia (Klotzsch) A.DC and Sphenanthera (Hassk.) Warb. (Smith 
et. al. 1986; Doorenbos et al. 1998). There are four endemic species, Begonia 
tribenensis C.R. Rao, Begonia minicarpa H. Hara, Begonia fiagellaris H. 
Hara and Begonia leptoptera H. Hara. 

Section Platycentrum (Klotzsch) A.DC. includes about 110 species 
from India through central China to Southeast Asia. and is one of the largest 
sections of the genus Begonia (Shui et al., 2002: Ye et al., 2004). This section 
is comprised of relatively large and robust species. characterised by a large 


144 Gard. Bull. Singapore 62 (1) 2010 


androecium on a distinct column, anthers with an extended connective, 
two highly convolute or spiralled styles and two-locular, drooping fruits 
with three unequal wings, one of which is considerably lengthened. 
During botanical expeditions from 2006-2008 to different parts of Nepal 
three new species of Begonia belonging to section Platycentrum were 
discovered, which are described and illustrated here, bringing the total 
number of Begonia species known from the country to 22. 


Species descriptions 


Begonia nuwakotensis S. Rajbhandary, sp. nov. [§ Platycentrum| 

Begoniae palmatae similis sed foliis basalibus longe petiolatis, laminae base 
cum lobis superpositis et in margine sine lobis vel dentibus longis curvatis, flore 
femineo tepalis exterioribus 4 similibus et tepalo interiore uno parvo, placentis 
trilamellatis et fructu cum ala laterali lata oblonga differt. —Typus: Central 
Nepal, Nuwakot, Kakani, Doman, 1700 m, 9 Aug 2007, S. Rajbhandary, S. 
Ranyitkar, K. P. Thapa and S. R. Bista S31 (holotype, E; isotype, KATH). Plate 1. 


Perennial, monoecious, rhizomatous, creeping herb, 14-30 cm tall. Stems 
rhizomatous at base, to 2 cm in diameter; erect portions semi-woody, reddish 
brown, becoming procumbent, tomentose, trichomes soft red. Stipules 
persistent, membranous, ovate, 16-21 x 8-10 mm, green to scariet, with an 
abaxially prominent midrib forming a thin, long appendage at the apex, 
margin entire, outer surface sparsely hairy on the midrib, trichomes red, 
inner glabrous and glossy. Leaves cauline, petiole 6-24 cm long, green, villose, 
trichomes, red, multicellular up to 2 mm long; lamina asymmetric, broadly 
ovate or oblate-orbicular, 5-22 x 5-19 cm, base slightly oblique, cordate, with 
overlapping lobes 1.5-3 cm long, margin remotely and shallowly denticulate, 
distinctly divided into short 0.5-2 cm long lobes, apex acute to acuminate; 
upper surface green strigulose/sericeous, trichomes small and white, lower 
surface pale green, densely pilose, trichomes red and long; palmately veined, 
7-8 veins. Inflorescences bisexual, axillary, cymose, few flowered, protandrous; 
peduncle 10-25 cm long, green, villose, trichomes red, multicellular. Bracts 
caducous, membranous, cymbiform, ovate, 12-15 x 8-11 mm, pale green to 
scarlet, glossy and pubescent, trichomes red, margin ciliate, apex acute. Male 
flowers: pedicel 1-2.3 cm, green, hairy, trichomes multicellular, red; tepals 4-5, 
white, outer 2 tepal orbicular, 13-15 x 13-20 mm, abaxially pilose, trichomes 
red, base cordate, margin ciliate, apex rounded to acute; inner 2-3 tepals 
obovate, 12-17 x 8-12 mm, smallest innermost 10-12 x 5-6 mm, white, 
glabrous, base cuneate, apex acute or sometimes rounded; androecium 
receptacle flat, free filaments almost equal, but slightly longer in the middle 
of the androecium, stamens ca 100, filaments 1-2 mm; anthers narrowly 


Three New Species of Begonia from Nepal 145 


Plate 1. Begonia nuwakotensis S. Rajbhandary. A. habit; B. Male flower; C. Stamens; D. 
Androecium; E. Upper leaf surface; F. Lower leaf surface; G. Bracts; H. Female flower; I. 
Stigma; J. Ovary; K. Prostrate stem; L. Fruits; M. Transverse section of fruit; N. Pollen; O. 
seed. 


146 Gard. Bull. Singapore 62 (1) 2010 


globose, on a small column, 6-10 mm diameter, yellow, symmetrical, obovoid, 
1.2-1.5 mm long, anthers towards the outside are longer than the filaments 
and those at the centre are shorter than the filaments, dehiscing through 
unilateral slits as long as the anther, connective slightly extended. Female 
flowers: pedicel 1-2 cm, hairy, trichomes red, multicellular; tepals 5, white, 
unequal, four outer tepals are orbicular, 20-22 x 18-22 mm, innermost tepal 
smallest, ovate, 13-15 x 7-8 mm, except for the innermost all the tepals are 
abaxially pubescent, margin entire, base rounded, apex rounded, acute in 
the innermost tepal; styles 2, deciduous, 3-6 mm long, fused at the base and 
ends into a broad wide apex, yellowish green; stigma green, a papillose 
band, strongly spiralled over the broad apex of the style and twisted at two 
ends; ovary green, pubescent, trichomes red, 2 locular; placentation axile, 
trilamellate. Fruits with pedicel 15-25 mm long, glabrous; 17-25 x 10-12 mm, 
nodding or pendulous, unequally 3-winged, with one larger wing and two 
narrow wings, larger wing oblong, 12-20 mm; lateral wings much smaller, 
5-6 mm, small wings recurved; capsule obovoid, ca 10-12 x 6-10 mm, green 
and hairy when young and dark brown when mature, glabrous; drying dark 
brown, dehiscing along the sutures between the two smaller wings. 


SEM studies: Pollen: large, perprolate with slightly concave sides and slightly 
pointed poles, irregular striate ornamentation with few scattered pores, P 
22-22.5 um, E 7.6-8.4 um, P/E 2.6-2.9, the endoporus elliptical with costae, 
margo distinct, narrow. Seeds: ellipsoidal to obovoid, mean size 325 x 240 
um; collar cells elongated, testa cells polygonal with straight or slightly 
curved anticlinal walls; anticlinal walls of testa cells thin and slightly raised; 
operculum short, flat and nipple-shaped; cuticular pattern consisting mainly 
of short linear zigzag striate ornamentation. 


Distribution: Known only from the type locality (Fig. 1). 
Ecology: Growing on wet shady slopes by the side of waterfalls, at ca 1700 m. 


Notes: This species is similar to Begonia palmata but differs in lacking long 
and curved lobes or teeth in the leaf margin. It also differs in having female 
flowers with 4 similar outer tepals and one small inner tepal, trilamellate 
placentae, and a fruit with a broad oblong wing. It is known only from 
the type locality and so it is named after the place of its collection. The 
population at the moment is healthy and reproducing well. However, as the 
single locality is not under protection and the capability of the species to 
cope with less shade and moisture is not likely to be good, we consider it to 
belong to the IUCN threat category VUD2. 


Three New Species of Begonia from Nepal 147 


Figure 1. Distribution of Begonia taligera, Begonia nuwakotensis, Begonia panchtharensis in 
Kaski, Nuwakot and Panchthar Districts, Nepal. 


Begonia panchtharensis S. Rajbhandary, sp. nov. [§ Platycentrum|] 
Begoniae sikkimensi similis sed foliis solum basalibus praesentis eis magnis 
suborbicularibus profunde lobatis longe petiolatis et fere glabris, floribus 
femineis 6-tepalis et fructu cum ala laterali lata falcata differt. -Typus: 
Cultivated plant grown from Eastern Nepal, Panchthar, Tinubote, Sisire, 
Prangbung, VDC, 2240-2300 m, 2 x 2007, U. Thamsuhang s.n., vouchered as 
S. Rajbhandary S74 (holotype, E, isotype, KATH). Plate 2. 


Perennial, monoecious, rhizomatous herb, 50-60 cm tall. Stem rhizomatous, 
8-12 cm long, 20-30 mm in diameter, covered with long roots. Stipules 
caducous, membranous, cymbiform, broadly ovate, 18-25 x 8-10 mm, pink 
with a light green tinge, glabrous with red spots on the abaxial surface, 
margin entire, apex acuminate. Leaves several, arising from the rhizome, 
petiole 33-46 cm long, succulent, glabrous, grooved, yellowish green with 
red linear dots; lamina slightly asymmetric, suborbicular and deeply lobed, 
24- 40 x 36-48 cm, base deeply cordate; margin irregularly serrulate or 
deeply dentate, distinctly divided to 2/3 of leaf length; lobes 6-8, apex 
acuminate, dark green above and pale green below, adaxially sparsely 
hirsute more so on the periphery, trichomes small, multi-cellular, crimson or 
whitish transparent, abaxially glabrous except for sparse hairs on the veins 
near the periphery, palmately veined, 6-8 veins. Inflorescences bisexual, 
axillary, cymose, dichotomously branched, protandrous; peduncle 42-50cm 
long, glabrous, grooved, sub-woody, yellowish green with red linear dots. 
Bracts caducous, in unequal pairs, membranous, cymbiform, elliptic to 
broadly elliptic, 25-30 x 15-18 mm, green, narrowly ridged abaxially and 
with linear or circular red granules, papery, margin entire, apex acuminate. 


148 Gard. Bull. Singapore 62 (1) 2010 


Plate 2. Begonia panchtharensis S. Rajbhandary. A. Habit; B. Male flower; C. Androecium; D. 
Stamens; E. Bracts; F. Rhizome; G. Petioles; H. Leaf venation; I. Leaf surface; J. & K. Inflorescence; 
L. Style and stigma; M. Stipule; N. Fruits; O. Transverse section of fruit; P. Seed; Q. Pollen. 


Three New Species of Begonia from Nepal 149 


Male flowers: pedicel 1-8 cm, light pink with linear red spots. glabrous: 
tepals 4, white to pink: outer 2 tepals broadly ovate. 1.2-1.8 x 0.9-1.2 cm, 
abaxially glabrous. with a pink tinge near the tip and the margin, base 
truncate, margin entire. apex acute: inner 2 tepals oval to narrowly oval. 
1.5-1.8 x 0.8-1.3 cm: white. glabrous, base cuneate, apex retuse: androecium 
globose, on a small column. 9-12 mm long. yellow. symmetrical. receptacle 
slightly raised: stamens 125-141. filaments 0.5-2.5 mm long: anthers oblong. 
1-1.5 mm, anthers towards the outside are longer than the filaments and 
those at the centre are shorter than the filaments, dehiscence through long 
straight slits which are slightly curved at the tip. connectives extended. acute. 
Female flowers: pedicel 1.5-2 cm. glabrous: tepals 6-8, outermost tepals pink. 
innermost white. symmetrical. oval 9-11 x 8-11 mm, glabrous. base rounded. 
Margin entire, apex acute: styles 2. persistent, 2.5-6 mm long, fused at the 
base. greenish yellow. stigmatic surface spirally convolute, papillose: ovary 
oblong and slightly curved, locular part 10-13 x 3-4 mm. glabrous with red 
- circular or linear granules. with three unequal wings: largest wing oblong 
with a rounded tip and two smaller more or less equal rim-like blunt wings: 
2-locular, placentae axile, bilamellate. Fruits: pedice] 2.5-3 cm, glabrous: 10- 
25 x 10-15 mm, nodding or pendant, falcate. unequally 3-winged: longer 
wing obovoid—oblong, 9-25 x 10-15 mm. lateral wings smaller, 1 mm long. 
wings red, capsule oblong, 10-15 x 2-4 mm. light greenish yellow when young 
becoming brownish green when mature, glabrous: drying dark brown with 
green tinge, dehiscing along the sutures between the two smaller wings. 


SEM studies: Pollen: large. perprolate with smooth convex sides and rounded 
poles, faint striate ornamentation with pores, P 16.8-21 um. E 8-10 um. P. 
E 2.1, the endoporus elliptical with costae. margo present. the grains not 
syncopate. Seeds: ellipsoidal, 325-345 x 215-240 um: collar cells elongated 
with straight and slightly curved anticlinal walls: testa cells polygonal with 
straight anticlinal walls; operculum long nipple-shaped with layers of cells: 
anticlinal boundaries broad and flat: the cuticle on the testa consisting 
mostly of patches without orientation while in the collar and operculum the 
cuticule has a long linear loose striate ornamentation. 


Distribution: Known only from the type locality (Fig. 1). 


Ecology: Growing on shady river banks and edge of the forest near rivers at 
ca. 2240-2300 m. 


Notes: This distinct species is somewhat similar to Begonia sikkimensis A. 
DC. in leaf shape but differs in having only basal. large and almost glabrous 
leaves, female flowers with six tepals, and broad falcate winged fruit. As it is 


150 Gard. Bull. Singapore 62 (1) 2010 


known only from the type locality, the species is named after the place of its 
collection. As the single locality is not under protection and the capability 
of the species to cope with less shade and moisture is not likely to be good, 
we consider it to belong to the IUCN threat category VUD2. 


Begonia taligera S. Rajbhandary, sp. nov. [§ Platycentrum|] 

Begoniae hatacoa affinis sed folii margine subintegro usque remote triangulari- 
serrulato dentibus interdum attenuatis, floribus haud lineatis, stigmate spiraliter 
torto, fructu cum ala laterali longa lataque apice rotundato et in petiolis 
radicibus ubi e planta separatis effractisve crescentibus recedit. —Typus: 
Central Nepal, Kaski, Bharat Pokhari, near Pokhara 700-740 m, 5 Sep 2007, 
S. Rajbhandary and § Adhikari S52 (holotype, E; isotype, KATH). Plate 3. 


Perennial, monoecious, caulescent herb, 30-70 cm tall. Stem rhizomatous 
at base, subwoody, 3.5 cm in diameter, covered with long roots; erect 
portions terete, 17-32 cm long, dark brownish red, branched, hairy. Stipules 
persistent, membranous, lanceolate, 6-11 x 7-13 mm, cymbiform, greenish 
white, transparent, abaxially glabrous, glossy, apex acuminate. Leaves 
cauline, petiole 6-10 (-32) cm long, succulent, densely covered with brown 
scales or multicellular two-armed short stalked trichomes, dark reddish 
brown at the base becoming ruby red adaxially near the top, readily forming 
adventitious roots when detached or broken; lamina asymmetric, cordate, 
ovate-lanceolate to elliptic-lanceolate, 10.5-23 x 5.5-18 cm, base slightly 
asymmetric, rounded or shallowly cordate, margin entire to attenuate- 
dentate, apex long-acuminate to attenuate, deep green adaxially, glossy, 
glabrous, yellowish-green abaxially, hairy with veins being covered densely 
with brown multicellular two-armed stalked trichomes, palmately veined, 
5-7 veins. Inflorescences bisexual, axillary, branched cymose, protandrous, 
peduncle glabrous 6-20 cm long, red, bearing few terminal flowers. 
Bracts: caducous, in unequal pairs, membranous, cymbiform, long elliptic 
to lanceolate, 5-8 x 1.5-3.5 mm, light green, transparent, apex acuminate, 
margin entire, glabrous, glossy. Male flowers: pedicel 1.5-3.5 cm, white to 
brownish green, glabrous; tepals 4, outer 2 tepals broadly oval, 9-17 x 7- 
12 mm, abaxially glabrous and glossy with a green or pinkish tinge more 
on the upper tepal, base rounded, margin entire, apex acute; inner 2 tepals 
oval, 9-18 x 7-10 mm, white, glabrous, base rounded, apex obtuse or retuse; 
androecium a loose sessile globose cluster, 7-9 mm long, golden yellow, 
symmetrical, receptacle slightly raised; stamens 90-95, filaments 1-2 mm long 
and free; anthers narrowly obovate, 2-2.5 mm, longer than the filaments, 
dehiscence through long straight slits, connectives extended, round or acute. 
Female flowers: pedicel 1-2 cm, glabrous; tepals 5, white, outer 2 tepals 
broadly ovate, 1-13 x 1-12 mm, glabrous, glossy, base rounded, margin entire, 


Three New Species of Begonia from Nepal 15] 


= 


Plate 3. Begonia taligera S. Rajbhandary. A. habit; B. Male flower; C. & D. Androecium: E. 
Stamens; F. Rhizome; G. Upper leaf surface; H. Lower leaf surface; I. Bracts: J. Broken petiole 
with adventitious roots; K. Female flower; L. Style and stigma; M. Two-armed trichome; N. 
Ovary; O. Transverse section of fruit; P. Pollen; Q. Fruit; R. Recurved small wings of the fruit; 
S. Seed; T. Androecium. 


152 Gard. Bull. Singapore 62 (1) 2010 


apex obtuse or rounded, the inner 3 oblong or lanceolate, 1.5-2 x 6-7 mm, 
apex obtuse to acute, but sometimes all the tepals are either broadly ovate 
or all oblong; styles 2, sometimes 3-4, persistent, 0.4-0.5 mm long, free, each 
style ending into a broad apex and connate at the base, green when young 
and becoming more yellow when mature; stigmas bifid, spirally tortuous 
and papillose all over; ovary oblong, 9-12 mm long, glabrous and glossy, 
unequally 3-winged, 2-locular, placentae axile, bilamellate. Fruits: pedicels 
1.2-2 cm long, glabrous; 19-25 x 9-15 mm, pendulous, with 3 unequal wings, 
the largest wing broadly falcate, 15-25 x 9-14 mm, apex rounded, the other 2 
wings smaller and narrow, ca 4-5 mm long with rounded tips, capsule oblong, 
10-15 x 3-4 mm, reddish brown when young and pale brown when mature, 
glabrous; drying pale brown, dehiscing along the sutures between the two 
smaller wings. 


SEM studies: Pollen: large, perprolate, pollen grain with smooth convex 
sides, rounded poles, compact irregular striate ornamentation with pores in- 
between, P 17.4-20 um, E 8 um, P/E 2.1-2.5, elliptical endoporus with costae, 
margo present around the endoporus, the grains are not syncolpate. Seeds: 
ellipsoidal, mean size 392 x 228 um; collar cells elongated with straight 
anticlinal walls; testa cells polygonal with straight and slightly curved 
anticlinal walls; operculum very short, flat nipple-shaped with a layer of flat 
elongated cells; anticlinal boundaries thin and slightly elevated; the cuticule 
with long linear striate ornamentation. 


Distribution: Known only from the type locality (Fig. 1). 
Ecology: Growing on moist shady rock ledges near streams at ca. 700-740 m. 


Notes: Allied to Begonia hatacoa Buch.-Ham. ex D. Don but differs in 
having an entire to attenuate-dentate leaf margin, while in B. hatacoa the 
leaf margin is usually always entire or only slightly dentate. Begonia taligera 
also lacks striped flowers, the stigma is spirally tortuous, the lateral long 
wing of the fruit is more broad with a rounded tip, and in the pollen the 
ornamentation of the testa cells is compact with irregular striate with pores 
in-between and margo present. One of the characteristic features of this 
species is the development of roots when a petiole is broken or detached 
from the main stem. The epithet taligera is derived from the Latin talea, 
and means producing cuttings for propagation. The population is currently 
healthy and reproducing well. However, as the species is restricted to a 
single locality, which is not under formal protection, and the capability of 
the species to cope with less shade and moisture is not likely to be good, we 
consider it to belong to the IUCN threat category VUD2. 


Three New Species of Begonia from Nepal 153 


Acknowledgement 


The authors thank the curators of BM, E, K, KATH, MICH and TUCH 
for allowing access to herbarium material; Dr. Mark Watson and Dr. Colin 
Pendry of the Royal Botanic Garden Edinburgh for their support and 
encouragement; the team of the Lower Kanchenjunga-Singalila Ridge 
CEPF Project, Eastern Nepal, for their extensive help in the collection of 
specimens, and to Dr. Robert Mill for translating the diagnoses into Latin. 
The Critical Ecosystem Partnership Fund (CEPF) USA, WWF (Nepal), 
University Grants Commission (UGC), and the M.L. MacIntyre Trust are 
thanked for providing financial support for this work. Finally, special thanks 
go to S. Ranjitkar, K. P. Thapa, S. Adhikari and the S. R. Bista for their help 
during field trips. 
References 


Doorenbos,J.,M.S.M.Sosefand J.J.F.E de Wilde. 1998. The sections of Begonia, 
including descriptions, key and species lists (Studies in Begoniaceae V1). 
Wageningen Agricultural University Papers 98(2): 1-266. 


Hara, H., W.T. Stearn and L.H.J. Williams. 1978. An Enumeration of the 
Flowering Plants of Nepal. Vol. 1. Natural History Museum, London. 


Press, J.R., K.K. Shrestha and D.A. Sutton. 2000. Annotated Checklist of the 
Flowering Plants of Nepal. Natural History Museum, London. 


Rajbhandary, S. and K.K. Shrestha. 2009. Begonia flaviflora H. Hara 
(Begoniaceae), New Record for Flora of Nepal. Journal of Japanese 
Botany 84(1): 16-18. 


Shui, Y.-M., C-I. Peng and C.-Y. Wu. 2002. Synopsis of the Chinese species 
of Begonia (Begoniaceae), with a reappraisal of sectional delimitation. 
Botanical Bulletin of Academia Sinica 43: 313-327. 


Smith, L.B., D.C. Wasshausen, J. Golding and C.E. Karegeannes. 1986. 
Begoniaceae. Part I: Illustrated Key, Part II: Illustrated Species List. 
Smithsonian Contributions to Botany 60. Smithsonian Institution Press, 
Washington. 


Ye, H.-G., E-G. Wang, Y.-S. Ye and C.-I Peng. 2004. Begonia coptidifolia 
(Begoniaceae), a new species from China. Botanical Bulletin of Academia 
Sinica 45: 259-266. 


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Gardens’ Bulletin Singapore 62 (1): 155-171. 2010 15 


in 


Genetic Variation of Populations of Scutellaria slametensis 
and S. discolor (Lamiaceae) on Gunung Slamet. 
Jawa Tengah (Indonesia) 


SUDARMONO AND B.J. CONN> 


’ Centre for Plant Conservation — Kebun Raya Bogor. Indonesian Institute of 
Sciences (LIPI). Jl. Ir. H. Juanda No. 13. Bogor, Indonesia. 
; Email: s_darmono@ yahoo.com 
National Herbarium of New South Wales. Mrs Macquaries Road. Sydney NSW 
2000, Australia. Email: barry.conn@rbgsyd.nsw.gov.au 


Abstract 


_ Genetic variation within and between populations of Scutellaria slametensis 
Sudarmono & B.J.Conn and S. discolor Colebr. on Gunung Slamet (Jawa 
Tengah, Indonesia) are evaluated by allozyme electrophoresis. Gels stained 
by 4 enzyme systems, namely, Aspartate aminotransferase (Aat). Esterase 
(Est). Malate dehydrogenase (Mdh) and Peroxidase (Per). were used to 
evaluate the number of polymorphic loci. The mean of total number of 
observed alleles per locus (A), mean of total number of effective alleles 
per locus (A-), percentage of polymorphic loci (Pp%). and expected genetic 
heterozygosity (H-) have been generated as parameters of genetic variation. 
The interpopulation genetic differentiation (Fsr) and estimated geographic 
distance between populations were used to evaluate the correlation 
between genetic differentiation and geographic effect. It was found that S. 
slametensis is genetically distinct from S. discolor (D = 1.4572). The mean 
genetic variation of S. slametensis (Pp = 75 %,A =2.00, Hz =0.450) is greater 
than that of S. discolor (Pp = 25 %, A = 1.25, He = 0.125). Almost all loci of 
the latter species are monomorphic and homozygotic. especially population 
9 near Baturaden (Pp = 0%: He= 0; Allele frequencies all = 1). There is 
a moderately high degree of variation between populations of these two 
species (Fst = 0.585, SE + 0.092). whereas within-population variation is 
low (1—Fsr= 0.415). Both species are out-breeding (at subpopulation level: 
Fis = -0.973, SE + 0.015: and population level: Fir = 0.180, SE + 0.183), with 
low levels of gene flow within and between populations (N= = 0.249. SE = 
0.065). The chromosome number of S. slametensis and S. discolor is 2n = 24. 


156 Gard. Bull. Singapore 62 (1) 2010 


Introduction 


Scutellaria (Lamiaceae) is the largest genus of the family with about 
360 species (Huang, 1994; Paton, 1990, 2004). The genus is widespread, 
subcosmopolitan, but poorly represented in moist tropical lowlands. There 
are currently four known species in Indonesia, namely, S. discolor Colebr., 
S. indica L., S. javanica Jungh. and S. slametensis Sudarmono & B.J.Conn 
(Backer and Backhuizen van den Brink Jr, 1965; Keng, 1978; Steenis van, 
1972: Sudarmono and Conn, 2010). The Indonesian species are all members 
of subgenus Scutellaria sect. Scutellaria (Paton, 1990) and are informally 
classified by Paton into several ‘species-groups:’ ‘S. discolor species-group’ 
(including S. discolor and probably S. slametensis);*S. humilis species-group’ 
(including S. javanica); *S. violacea species-group’ (including S. indica). 

Allozymes have provided the most abundant source of information 
regarding genetic variation in natural populations (Hamrick and Godt, 1990). 
Genetic variation parameters such as amount of interbreeding between 
populations, allele heterozygosity, genetic diversity, genetic differentiation 
and the amount of gene flow can indicate the level of gene mutation, genetic 
drift, genetic ‘bottle necks’, and even the possible level of vulnerability of 
endangered populations. These parameters are equally useful for assessing 
genetic variation within and between populations of a species as they are 
between species (Hamrick and Godt, 1990). Maki (1972) and Luzuko et al. 
(2000) found that there was a significant correlation between low gene flow 
and geographic distance or isolation by distance. Wright (1943) was the first 
to describe the genetic process of isolation by distance that may operate 
when populations conform to a ‘stepping stone’ model, where gene flow 
occurs only between adjacent populations. He demonstrated that random 
genetic drift within localized populations, combined with limited migration 
among populations, can result in increased genetic differentiation with 
increasing spatial distance between populations. 

Grant (1981) concluded that geographic isolation was an important 
factor that may result in allopatric speciation with reproductive isolation 
playing an important role in sympatric speciation. Both of these speciation 
phenomena might occur within populations of Scutellaria on Gunung Slamet. 
Scutellaria slametensis occurs at elevations of more than 1,000 metres, 
whereas S. discolor occurs at elevations of less than 800 m. Chromosome 
numbers were recorded and within-population allozyme variation was 
analyzed for both species. 


Genetic Variation of Populations of Scutellaria slametensis lay) 


Materials and methods 


Sample collections 

Samples of Scutellaria slametensis were collected from six populations on 
Gunung Slamet (Jawa Tengah, Indonesia, Fig. 1). These populations occur 
at different elevations. Populations | (elevation 1,980 m) and 3 (1,802 m) 
occurring on the southwestern slopes, along the Kaligua and Brebes route 
in the Perseroan Terbatas Perkebunan Nusantara (PTPN) IX National 
Tea Plantation (in Paron and Sokarata Blocks of the Protected Forest, 
respectively), Forest Holding Unity Division — Balai Kesatuan Pemangku 
Hutan (BKPH), with population 2 occurring closer to the summit of Gunung 
Slamet (at 2,002 m) (Table 1). Populations 4-6, from the southern part of 
Gunung Slamet, on the Baturaden climbing track occur between elevations 
of 1,390 and 2,215 m (Table 1), on the eastern and southeastern slopes. 
Samples of three populations of S. discolor were collected at elevations less 
than 800 m, from the Baturaden area of Gunung Slamet (Table 1: populations 
7-9). Although Keng (1978) concluded that S. discolor occurred over a wide 
range of elevations, from 500-2,400 m, this species was not found above 
800 m. on Gunung Slamet. Ten individual plants were sampled from each 
population. Collections of all populations of both species were cultivated 
at Kebun Raya Bogor (Indonesia) and dried herbarium vouchers lodged 
at Herbarium Bogoriense (BO), Herbarium of Kebun Raya Bogor (KRB) 
and National Herbarium of New South Wales (NSW). 


4 Baturaden 


eoreters 


Figure 1. Distribution map of populations of Scwtellaria included in this study on Gunung 
Slamet, Jawa Tengah, Indonesia. Populations 1-6 = S. slametensis; 7-9 = S. discolor, red square 
= summit of Gunung Slamet, elevation 3,428m. 


158 Gard. Bull. Singapore 62 (1) 2010 


Chromosome observation 

Growing root tips were incubated in 0.05 % colchicine aqueous solution 
for 2 hours at 18°C. They were fixed with the fixative solution (ethanol: 
chloroform:glacial acetic acid = 2:1:1) for more than 45 minutes at 5°C. The 
root tips were then macerated with 1N HCl at 60°C for 18 seconds. The 
meristematic tissues were stained with 2% aceto-orcein for 5-10 minutes on 
a glass slide, one drop of 45% acetic acid was added and the tissue covered 
and squashed gently. 


Allozyme analysis 

Within-population allozyme variation was analyzed by enzyme systems 
for six populations of Scutellaria slametensis and compared to the within- 
population variation found within three populations of S. discolor, both 
occurring on Gunung Slamet (Jawa Tengah, Indonesia, Figure 1). 

Leaf samples were prepared for allozyme electrophoresis by extracting 
0.5 cm’ samples of fresh leaf material. Young leaves were ground with 0.1 M 
Tris-HCl pH7.5,0.1% 2-mercaptoethanol,0.001 M EDTA (tetrasodium salt). 
Extract was absorbed by filter paper (Whatmann No.3) andrun by 12% starch 
gel (4.5 hours, 300 volt; 45 mA). Four horizontal enzyme systems analyzed 
were Aspartate aminotransferase (Aart; EC 2.6.1.1), Esterase (Est; EC 3.1.1), 
Malate dehydrogenase (Mdh; EC 1.1.1.37) and Peroxidase (Per; EC 1.11.1.7). 
Staining procedure followed Soltis et al. (1983), with some modification in 
buffer pH and concentration (Mdh buffer with 1.5 M TRIS-HCI pH 8.0). 
The locus specifying the most anodally migrating isozyme was designated 
as 1, the next 2, and so on. Similarly, the most anodally of a gene was labeled 
‘a’, the next ‘b’ and so on, as done by Shield er al. (1983) and Kephart (1990). 

The genetic variation of populations is expressed as a percentage 
of polymorphic loci (Pp %), mean number of observed alleles per locus 
(A), and Nei’s unbiased genetic diversity or expected heterozygocity (/7.), 
using POPGENE version 1.32 (Yeh et al., 1999). Gene flow (Nm — the 
number of migrations per generation) for all loci was also estimated. The 
analysis of genetic identities (7) and genetic distance (D) for each pair-wise 
combination of populations were also estimated following Nei (1978). In 
this study, unbiased genetic identity was used to accommodate for the bias 
caused by small sample size (<50 individuals). 

There are several indices used to evaluate genetic diversity, including 
total genetic diversity (Hr), genetic diversity within populations (Hs), 
genetic diversity among populations (Ds7), and the among populations 
genetic differentiation coefficient (Gsr). These statistics were generated 
using FSTAT program (Goudet, 2002). Total genetic diversity (7/7) was 
partitioned into within-population (Hs) and between-population (Dsr) 
components. Hr was estimated for each polymorphic locus from following 


Genetic Variation of Populations of Scutellaria slametensis 159 


equations: 
Hr =Hs+ Dsr 


oP) 
v= 5 


Where, ji is gene identity in sub-population and S is number of sub- 
populations. ji was estimated by: 


ji = Sa 
Where, xis the frequency of the Ath allele in ith sub-population 


Genetic diversity between sub-populations (Ds) was estimated by: 
Dsr= ( Ly ID¥), Si 


1 
Dy= ) 5 Pa-Pa¥ 


Where, Pix is the frequency of the kth allele in ith sub-population, while Pi 
is frequency of kth allele in jth sub-population. 


Gene differentiation that occurs among populations (Gsr) was examined 
with Nei’s genetic diversity indices (Nei, 1977, 1986). Gsr was expressed 
relative to total genetic variations among populations as: 


Dsr 
Gsr = 


Mean values of Hr, Hs, Dsr and Gsrwere the average of all polymorphic 
loci within each group. All calculations were generated for each pair-wise 
comparison of populations and species using FSTAT program (Goudet, 
2002). 


The genetic structure of the studied populations was also analyzed in term of 
the following F-statistics: Fis - fixation index related to non-random mating 
within populations; Fir - mean inbreeding coefficient, Fs7 - interpopulation 
genetic differentiation (following Weir and Cockerham, 1984). 


Gene flow (N,,) for all loci was estimated: 


_G~6y) 


N, 
m 4Gsr 


160 Gard. Bull. Singapore 62 (1) 2010 


Assuming that populations have reached equilibrium between the effects 
of migration and random genetic drift, the degree of population subdivision 
was quantified using: 


1 
Fy => 
st” (4N,, + 1) 


Fsr is the proportion of total genetic variance contained in subpopulation 
S relative to total genetic variance 7. Unbiased data matrices (Nei, 1986) 
were generated using POPGENE (Yeh et al. 1999). It was also used to 
calculate mantel test (Sokal and Rohlf, 1995) for testing the null hypothesis 
of independence between genetic differentiation and geographic distances 
separating populations (Yeh et al., 1999). Allozyme data were analyzed 
using UPGMA (Unweighted Pair-Group Method using Arithmetic 
Average) clustering techniques to construct a dendrogram to assist in the 
interpretation of these genetic data between species and populations. The 
dendrograms were generated using NTS YS (Rohlf, 2000). 


Results 


Chromosome analysis 

Chromosome number of Scutellaria slametensis (Figs.2A & B) and S. discolor 
(Figs. 2C & D) are both diploid 2n = 24. The length of mitotic metaphase 
chromosome of S. slametensis varies from 2-2.5 um, whereas those of S. 
discolor are smaller (1.5-2 um). 


Figure 2. Mitotic metaphase chromosomes of Scutellaria slametensis (A, B) and S. discolor 
(C, D), both 2n=24. A & C, microphotographs; B & D, line-drawings of microphotographs. 
Scale bar = 5 um. 


Genetic Variation of Populations of Scutellaria slametensis 161 


Genetic Variation 

Four loci were detected in all populations of both species of Scutellaria 
(Table 1). The four enzyme systems examined (namely, Aat, Est, Mdh and 
Per) were consistent for all nine populations studied. Populations 1, 4-6 
(Table 1) are genetically highly variable with 75% of loci polymorphic (Pp). 
Population 4 has the highest mean number of alleles per locus (A = 2.00), 
with a high mean number of A=1.75 for populations 1,5, 6 (Table 1). The 
expected heterozygosity is also regarded as high for these four populations 
(population 1: He= 0.375; 4: He = 0.401; 5: He= 0.375; 6: He = 0.375) (Table 
1). In contrast, the lowest genetic variation occurs in population 9 (Pp= 0 
%,A = 1.00, He = 0.000) (Table 1). Others populations that are regarded as 
having low genetic variation include: population 2 (Pp = 25 %, A=1.25, He= 
0.139) and population 8 (Pp = 25 %, A = 1.25, He = 0.125). 


Table 1. Mean genetic variation of populations of Scutellaria slametensis and S. discolor 
on Gunung Slamet (Jawa Tengah, Indonesia). Ten samples of each population (N) were 
examined. Populations 1-6 = Scutellaria slametensis; 7-9 = S. discolor. Percentage polymorphic 
loci (Pp), mean number of observed alleles per locus (A), mean of effective alleles per locus 
(Ae), mean of expected heterozygosity (He) (Nei, 1978). *Comparison of genetic variation 
categorized at regional geographic distributed means Pp=36.4 %, A=1.55, Ae =1.16 and He 
= 0.118 (Hamrick and Godt, 1990). 


Eep Location eititnde Geocode Pp% A Ae He Category * 
no. (m) 
1 Protected forest block 7.2683° S z 2 2 ; 
: : : fo Ths) 1S: 75 0.375 high 
Paron, Kaliwadas oe 109.1584° E Te : 's 
2 __ Protected forest block, A pe SUIS ae - z 
, 2,002 ae 25 1255 125) 05139) Sbow 
Sakub, Kaligua a 109.1456° E 
3 Protected forest block 7. 2829° S 2 P E e 
: , 1,802 z 50 =1.50 1.50 0.250 Low 
Sokarata, Kaligua 109.1298° E 
4 Protected forest, Petak II 1,390 eS 75 2.00 1.88 0.401 high 
2 s 109.2004° E a = i 
5 Protected forest, Post III, 2215 7.2668° S 75 1.75. 1.75 0.375 high 
Baturaden 109.2011° E 
6 Protected forest, Post I 7.2668° S : = ae =e 
z 3 5778 Lee 75 1.75 1.75 0.375 high 
Baturaden 109.2023° E 
7 Camping ground, 7.3119°S = Bs Z = 
rie 771 ore 50 1.50 1.50 0.250 Low 
Baturaden 109.2351° E 
| 30522 S egy er ee 
8 Baturaden Botanic Garden 771 109.2333°E 1.25 1.25 0.125 Low 
7.3069°S 


9  Telaga Sunyi, Baturaden at 0 1.00 1.00 0.000 low 


109.2427° E 


162 Gard. Bull. Singapore 62 (1) 2010 


Population structure and gene flow 

Genetic diversity indices (Table 2) demonstrate that Aat-] and Mdh 
contain the highest total diversity (Hr) of those loci surveyed with 62.1% 
and 64.8%, respectively. The enzyme Est-1 contained the least genetic 
diversity (Hr= 44.4%). The mean of total genetic diversity (H7) was 58% 
(SE +0.030), whereas genetic diversity within populations (Hs) was 26.7% 
(SE +0.058) and among populations (Dsr) 31.3% (SE +0.034). There was no 
genetic diversity within-populations in the Est-/ locus (Hs= 0%). Likewise, 
the genetic diversity within and among populations of Aat-/ was low (Hs= 
29.1% and Dsr = 35.1%, respectively). Although the genetic diversity within- 
populations of Mdh and Per were similar (both Hs = 38.9%), the among 
population genetic diversity was Dsr= 25.9% and Dsr= 19.8%, respectively. 
The mean genetic differentiation between populations (Gs7) is 57.1% (SE 
+ 0.095). 


Table 2. Genetic diversity indices, F-statistic and estimation of gene flow between populations 
of Scutellaria. Hs, the genetic diversity within populations; Dsr, the genetic diversity among 
populations; Hr, the total genetic diversity; Gsr, the among populations gene differentiation 
coefficient; Fis, the fixation index related to non random mating within populations; Fir, 
the mean inbreeding coefficient of a set of a populations; Fs7, the interpopulation genetic 
differentiation; Nm, gene flow estimated from Fsr = 0.25(1 - Fsr)/Fsr. SE, standard error. 


Genetic diversity indices F-statistic Gene flow 
LOGTS Hs  Dsr Hr Gsr Fis Fir Fsr Nm 
Aat-1 0.291 0.351 0.641 0.547 | -0.919 0.135 0.549 | 0.205. 
Est-1 0.000 0.444 0.444 = 1.000 ht! 1.000 1.000 0.000 
Mdh 0.389 0.259 0.648 0.400 | -1.000 -0.091 0.455 0.300 
Per 0.389 0.198 0.587 01337 | -1.000 -0.326 0.337 0.492 
Mean 0.267 0.313 0.580 0.571 | -0.973 0.180 0.585 0.249 
SE 0.058 0.034 0.030 0.095 | 0.015 0.183 0.092 0.065 


The estimation of the genetic variation among and within populations 
indicates a moderately high degree of differentiation among populations 
(Fst = 0.585; SE +0.092), whereas variation within-populations (1-Fs7) 
represented 41.5% of the total variance. These species exhibit a considerable 
degree of out-breeding at both the subpopulation (Fis = -0.973; SE +0.015) 
and population levels (Fir = 0.180; SE +0.183), even though the estimate 
of mean gene flow was low (Nm = 0.249; SE +0.065). However, the null 
hypothesis that these populations are at equilibrium is rejected because 
there is no correlation between pair-wise genetic differentiation values (Fs7) 
and geographic distance (r = 0.365; Mantel t-test= 2.259; p < 0.01) (Fig. 4). 


Genetic Variation of Populations of Scutellaria slametensis 163 


Based on genetic distance (Nei 1978), two population-groups are 
differentiated (Fig. 3), representing Scutellaria discolor (populations 7-9) 
which is genetically distinct from all populations of S. slametensis (Genetic 
distance, D=1.4572). Within the populations of the latter species, populations 
1 (Paron block, Brebes) and 5 (population in Post III, along walking track 
from base camp Baturaden to summit of G. Slamet) are genetically similar 
(Fig. 3). Likewise, populations 4 (Post II) and 6 (Post I) are also genetically 
similar and this latter population pair is genetically close to the previous 
pair. Populations 8 and 9 (S. discolor) are also genetically similar (Fig. 3). 


Allele frequency of 9 populations and allele shared among species 

Most alleles are relatively common within the sampled populations 
(occurring in at least 50% of loci) (Table 3). However, two alleles with low 
frequencies in population 4 are Aat-1 with only ahs of chromosomes carry 
this allele and Aar-I" (35%). Alleles Aat-1 , Mdh-I' and Per’ were shared 
between S. slametensis and S. discolor, whereas, Aat-1° ’, Est-I° and Per" were 
specific to S. slametensis. Alleles Est-1 and Per are specific to S. discolor. 


Table 3. Allele frequency of 4 polymorphic loci in 9 populations of Scutellaria. Populations 
1-6 = S. slametensis; 7-9 = S.discolor 


Population Aat-I’ Aat-I’ Aat-I' Est-I° Est-I’ Mdh-I'  Mdh-I’ Mdh-I' Per" — Per’ Per’ 


1 0.50 0.50 1.00 0.50 0.50 0.50 0.50 
2 1.00 1.00 1.00 0.50 0.50 

3 0.50 0.50 1.00 0.50 0.50 1.00 

4 0.50 035 0.15 0.50 0.50 0.50 0.50 

5 0.50 0.50 1.00 0.50 0.50 0.50 0.50 

6 0.50 0.50 1.00 0.50 0.50 0.50 0.50 

7 1.00 1.00 0.50 0.50 0.50 0.50 
8 1.00 1.00 1.00 0.50 0.50 
9 1.00 1.00 1.00 1.00 

Discussion 


Paton (2004) noted that the chromosome numbers for Scutellaria are 2n=12- 
88, with most frequent numbers being 2n=20, 22, 24, 32 and 34. Chromosome 
numbers of Scutellaria subgenus Scutellaria, which includes all Indonesian 
species, are 2n=24-34 (Paton, 1990). Since the chromosome numbers of S. 
discolor and S. slametensis are both 2n=24, these two species are possibly 
closely related. Both of these species have small chromosomes, less than 5 
um long. 

Allozymes have been successfully used to compare mating system, 
migration and local differentiation within and between populations (Brown, 


164 Gard. Bull. Singapore 62 (1) 2010 


1990). Factors such as, regional distribution of a taxon, geographic range, 
breeding system, seed dispersal mechanisms and successional status 
have been associated with differences in the percentages of polymorphic 
loci (Pp%), mean number of observed alleles per locus (A), and genetic 
diversity within-populations (He) (Hamrick and Godt, 1990). Values 
of Pp%, A and He obtained for S. discolor and S. slametensis are very 
similar to those obtained for out-breeding plants, wide-spread species, 
and for those with seeds that are dispersed by gravitational forces. 

Scutellaria slametensis is a species with a restricted distribution and 
has mean genetic variation (Pp= 75 %, A = 2.00, He = 0.450) equivalent to 
that of S. montana (Pp = 75.42 %, A = 2.21, He = 0.19; see Cruzan, 2001), 
a species restricted to parts of Georgia and Tennessee (U.S.A.) (Cruzan, 
2001). Scutellaria slametensis exhibits greater genetic variation than plants 
of S. discolor sampled from G. Slamet. Almost all loci of the latter species 
are monomorphic and homozygotic, especially population 9 (Pp = 0 %; He 
= (); Allele frequencies all=1). 

The value of allozyme electrophoresis in delimiting taxa has been 
found to be useful by several researchers (such as, Coates and Hnatiuk, 
1990; Crawford, 1985; Gottleib, 1984). Although all populations of S. discolor 
and S. slametensis are relatively close geographically, they are genetically 
distant based on allozyme divergence. This is possibly a result of the steep 
mountainous terrain restricting the potential pollinators tosmall geographical 
areas within the region (Tyler, 2003). Within S. s/ametensis, populations 4 
and 6 are genetically and geographically close. Population 5 is unexpectedly 
genetically close to population | even though the two are not geographically 
close. Together, populations | and 5, and 4 and 6 form a genetically distinct 
grouping (Fig. 3). Although the two high altitude populations, 5 (elevation 
at 2,215 m) and 2 (2,002 m) are geographically relatively close, these two 
populations are genetically distant (Fig. 3). Although population 1 is located 
in between populations 2 and 3, the genetic variation, hence gene flow, 
within the latter two populations is low suggesting that they do not share 
a common population of pollinators. Contrary to this, within-population 
variation in the population group | and 4-6 is high, representing high 
gene flow between these populations (Table 1). The three populations of 
S. discolor (populations 7-9) are both geographically close and genetically 
similar. Species characterized by a low level of gene-flow, such as S. discolor 
(Table 1), or a low level within or between populations, such as populations 
2 and 3 (S. slametensis), may indicate high levels of selfing. Cruzan (2001) 
hypothesized that the smaller fragmentation threshold may reflect higher 
levels of selfing in isolated populations because of the absence of pollinators. 
Unpublished results from observations of floral behaviour and crossing 
tests of S. slametensis clearly demonstrate that this species is mostly self- 


— 


Genetic Variation of Populations of Scutellaria slametensis 165 


compatible as has been found in S. indica (Sun, 1999). However, within the 
current study, the low overall proportion of total genetic variance within a 
subpopulation relative to total genetic variance (Fsr= -0.9769) implies a low 
level of differentiation among populations, suggesting that S. discolor and S. 
slametensis are out-breeding. The low level of gene flow (mean Nm = 0.1988) 
observed suggests that although these populations are geographically close, 
they are relatively isolated genetically. Slatkin (1987) concluded that values 
of N» < 1 mean that genetic drift will result in substantial differentiation 
between populations. This is possibly a consequence of pollinators only 
visiting flowers within a restricted area, hence mostly visiting plants in 
closely adjacent areas (such as, between populations 4 and 6, and between 8 
and 9). However, this does not explain the lack of genetic distance between 
populations 1 and 5 (D = 0.000). 


] 

5 

4 

6 S. slametensis 
3 

2 

: S. discolor 

8 

9 

0.02 0.35 0.71 1.08 1.45 


Figure 3. Dendrogram of Nei’s genetic distance (Nei, 1978) between six populations of 
Scutellaria slametensis and three populations of S. discolor on Gunung Slamet, Jawa Tengah, 
Indonesia. 


Mean total genetic diversity (Hr) showed majority of partitioning 
occurred among populations (Dsr= 31.5%) rather than within populations 
(Hs = 0.267). The large amount of genetic differentiation among populations 


166 Gard. Bull. Singapore 62 (1) 2010 


(Gsr = 0.571; Fsr = 0.585) indicate strong genetic differentiation between 
these populations (Table 3). F-statistics have similar values and indicate that 
populations of S. discolor and S. slametensis are genetically structured (Table 
2). High levels of genetic structure within-populations are supported by the 
Est-I locus but with low levels of gene flow (Nm) (Table 2). High levels 
of pollen transfer indicate movement of pollinators that will naturally lead 
to increased genetic differentiation but decreased gene flow (Ellstrand and 
Elam, 1993). If genetic variation within species is predominantly affected 
by shared alleles, then there may be a significant correlation between 
shared alleles and the local geographical distribution pattern of the species. 
An understanding of nature of shared allele may be used to understand 
conservation implication for future. 

Throughout this study we have assumed a ‘stepping-stone’ model 
(Kimura, 1953) of population structure among these plants whose dispersal 
ability is constrained by distance such that gene flow is most likely to occur 
between neighbouring populations (Hutchinson and Templeton, 1999). 
Consequently, it would be expected that adjacent populations tend to be 
more genetically similar than more distant populations. Therefore, assuming 
a‘stepping-stone’ model of regional population structure, the null hypothesis 
that these regional populations are at equilibrium is rejected because there 
is no association between pair-wise genetic differentiation values (Fs7) 
and geographic distance (r = 0.365; Mantel t-test= 2.259; p < 0.01) (Fig. 4). 
In this study, the populations of Scutellaria on Gunung Slamet consist of 
mostly small, more or less isolated populations, such that allele frequency 
drifts independently of geographic distance between populations and is 
much more influential in determining the population structure than gene 
flow. These results are consistent with limited pollination events occurring 
between populations and/or the inhibition of dispersal in this mountainous 
terrain. 

The dendrogram of the sampled populations (UPGMA - based on Ne1’s 
genetic distances; Nei, 1978) indicates a correlation between shared alleles and 
genetic distance (D) (Fig. 3). The topology of this dendrogram is congruent 
with morphological differences between S. discolor and S. slametensis. Based 
on allozyme data, the greatest genetic distance (D =1.45) was found between 
S. discolor and S. slametensis. There are several morphological features that 
distinguish S. discolor and S. slametensis (Sudarmono and Conn, 2010). The 
shape of the leaf, height of stem and number of flowers at inflorescence 
nodes of S. discolor are very different from those of S. slametensis. Scutellaria 
discolor has flowers with corolla purple, whereas, S. slametensis has corollas 
that are white basally, pink distally (populations 1 & 2) or purple distally 
(populations 3-6). Scutellaria slametensis has obovate leaves, whereas 
those of S. discolor are ovate. The former species has two flowers at each 
node of the inflorescence, whereas S. discolor has four flowers per node. 


Genetic Variation of Populations of Scutellana slametensis 167 


0.63 
3 
S 0.47 
= 
: 
ps 0.32 
= 
2 0.16 
E . 
0.00 AE 5.45 8.18 10.90 
Geographic distance (km) 


Figure 4. Correlation between geographic distance (km) and genetic differentiation (Fs7) 
of nine populations of Scutellaria discolor and S. slametensis on Gunung Slamet. r=0.365: 
Mantel f test = 2.259: p < 0.01. 


Conclusion 


Scutellaria discolor and S. slametensis are morphologically and genetically 
distinct. Genetic variation within populations 1, and 4-6 are high (S. 
slametensis), whereas within populations 2, 3 (both S. slametensis), and 7-9 
(S. discolor) are low. Although the genetic diversity within S. slametensis 
is partitioned, with one group with high genetic variation (populations 1, 
4-6) and one with low values (populations 2 and 3). overall this species has 
a high level of genetic heterozygosity that makes it a species with a high 
frequency of heterogamy. Consequently, from a conservation point of view, 
this species is probably a low-risk species (Fracaro and Echeverrigaray. 
2006). Even though the two species at the subpopulation level (Fis = -0.973: 
SE +0.015) and population level (Fir = 0.180; SE +0.183) are out-breeding, 
the mean gene flow was low (Nm = 0.249; SE +0.065). Historically, genetic 
drift has affected the overall structure of the populations of this species, 
rather than gene flow. However, within the distribution of S. slametensis, the 
mean gene flow was high for the local populations 1. 4-6. Since this species 
is only known by a limited number of individuals in a restricted locality, it 
may prove to be vulnerable (IUCN, 2001) should the populations become 
more isolated, resulting in the level of gene flow and hence, heterozygosity 
becoming further reduced. 


168 Gard. Bull. Singapore 62 (1) 2010 


Although S. discolor is a widespread species, all loci of the plants 
sampled on Gunung Slamet are monomorphic and homozygotic. An 
assessment of the genetic variability of this species throughout its range 
is required. Even though a species may be widespread, an increased 
homozygosity may result in a reduction of the vigor of individuals, expression 
of deleterious characters, increased seed abortion, reduced fertilization and 
germination rates. These are factors that may lead to the disappearance of 
the populations (Dubash and Fenster, 2000). 


Acknowledgements 


The authors are much indebted to P. Daryono, P. Budiyanto, and P. Kartam 
who provided field assistance in the Protected Forest of Gunung Slamet from 
Kaligua, Kaliwadas and Baturaden. We thank Ir. Sinung Nugroho (BKPH 
Gunung Slamet Barat), Dr. Irawati (KRB) for permission to undertake 
this study, and Prof. Mien A. Rifai (BO) for comment on this manuscript. 
Prof. Dr. Syamsuardi, (Andalas University, Sumatera Barat, Indonesia) for 
critical comment on technical analysis of allozymes. The first author received 
financial assistance from DIPA-LIPI Pelaksanaan Kegiatan Penelitian Hasil 
Seleksi Workshop Kiat-Kiat Memenangkan Proyek TA 2008. 


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Gardens’ Bulletin Singapore 62 (1): 173-176. 2010 173 


A New Species of Polyalthia (Annonaceae) from Sabah 


I.M. TURNER 
Research Associate, Singapore Botanic Gardens, and 
Royal Botanic Gardens, Kew 


Abstract 


Polyalthia lasioclada 1.M. Turner, sp. nov. is described. It is a small tree 
known from the Mount Kinabalu area of Sabah. 


When looking through some Annonaceae specimens recently collected from 
the Kinabalu Park area of Sabah, I realised several were the same thing as 
an odd specimen I had tentatively placed in Polyalthia microtus Miq. The 
leaves drying greyish with auriculate bases and orange-red corollas clearly 
pointed to membership of the informal ‘Polyalthia insignis species-group’ 
of Johnson and Murray (1999). The unusual feature of the specimens was 
the villose twigs reminiscent of Polyalthia borneensis Merr. and P. bullata 
King rather than typical P. microtus. The availability of more collections 
confirmed that there were some consistent differences in flower and fruit 
form that separated the shaggy-twigged plant and therefore I here describe 
it as a new species. 


Polyalthia lasioclada \.M. Turner, sp. nov. 

Polyalthiae microtus ramulis_ petiolisque_ villosis, sepalis minoribus, 
monocarpiis globosis vix cylindricis, in sicco aureobrunneis laevibus vix 
brunneis verruculosisque differt. -Typus: Malaysia. Borneo, Sabah, Ranau 
District, west of Kg Takutan, 2500 ft, 25 May 1973, G. Shea & Aban, SAN 
77174 (holotype, SING; isotypes, K, L). Plate 1. 


Small tree. Twigs drying dark grey or grey-brown with shallow longitudinal 
grooves and some cracking, youngest parts brown or reddish brown densely 
covered with long (3-4 mm) more or less straight golden brown hairs. Leaves 
chartaceous to subcoriaceous, drying grey or grey-brown above, brown 
beneath, glabrous except for long straight hairs on midrib below, densest near 
base, midrib and lateral nerves immersed above, prominent beneath, lamina 
elliptic to elliptic-obovate, 4.5-20 x 1.5-7.7 cm, apex acute to shortly acuminate, 
base auriculate, lateral veins 15-19 pairs, angled or arching forward, looping 
distinctly via a distinct intramarginal, tertiary venation reticulate. Petioles 1- 


174 


Plate 1. Photograph of the holotype of Polyalthia lasioclada 1.M. Turner, sp. nov. 


Gard. Bull. Singapore 62 (1) 2010 


FLORA OF SABAH 
HERBARIUM OF TILE FORPSS VEYARTMENT SANDARAN 


No Sas TUT 
Family rt 
Botanical Nam= es met dasymathaly 


Verzacular 

Locality (sce Rerer F S of Gq. Tadeter, 

District Dare Al 2g Fu ust 
Collett gy gyne & Abas Date 24,57 he : 


Habitat snl} tres in asiecstery on stomp bilites - 
prises frre. 


Notss 
Welgiety 10 Pt, Sirthy 5 dace. 
ark; ditt ceay/ nem Clement, 
Teaver’ “icrore;red inh Coungte 
, / 
v J 
Dapl ‘ 0/PNH 
cociving deplicates ace eted wo cHmmunicais ew 
to the Conservator 
Gr > CW? 72 


A new species of Polyalthia 175 


5 mm long, 1-3 mm thick, densely villose. Inflorescences single-flowered, 
intranodal. Flowers with pedicel 12-24 mm long, less than 1 mm wide, 
drying red brown with scattered pale hairs, faintly wrinkled longitudinally, 
sepals ovate or triangular 1-2 x 2-3 mm, drying brown with pale hairs 
externally, glabrous within, petals lanceolate 15-30 mm long, orange-red, 
outer slightly wider than inner, 5 vs 3-4 mm, drying dark brown or almost 
black, outside slightly wrinkled longitudinally, minutely pimpled near the 
base with sparsely scattered short white hairs, inside smoother with hairs 
confined to the apex. stamens many ca 2 mm long, connective apex truncate, 
carpels ca 12, ca 1.5 mm long, densely pale hairs. Fruits with pedicel ca 18 
mm long, 1 mm wide, drying reddish brown, monocarps ca 12, globose, 8- 
10 mm diameter, apex apiculate, drying golden brown, sparsely adpressed 
pale hairy, drying relatively smooth, stipe 5-8 mm long, | mm thick. Seeds 1-2, 
drying pale brown. 


Specimens seen: MALAYSIA. Sabah, Kota Marudi District, Kampung 
Monggis, 4 batu dari pusat Kampung Monggis Utara, 9 Apr 1996, Matamin 
Rumutom 186 (K); Kampung Kawasan Taman 100 m dari Sungai Mokodon, 
8 Mar 1996, Daim Andau 344 (K); Kampung Kawasan Taman, Jalan ke Palu 
Agayo, 16 May 1995, Kinsum Bakia 44] (K); Ranau District, Kampung 
Nalumad 2 batu from Kg Nalumad, 3 Oct 1998, Daim Andau 1020 (K); 
Ranau District, Kampung Nalumad 5 batu dari Kg Nalumad, 9 Oct 1996, 
Daim Andau 855 (K). 


Notes: The chosen specific epithet is derived from Greek (/asios = shaggy, 
woolly, clados = branch, shoot) and reflects the characteristic villose twigs 
of the species. 

Polyalthia lasioclada is similar to the widespread and rather 
variable Polyalthia microtus, particularly in flower form. The sepals of P. 
lasioclada are generally smaller than those of P. microtus (1-2 x 2-3 mm 
versus 4-6 x 4-6 mm). The monocarps of P. lasioclada are dry globose, 
relatively smooth and a golden brown, whereas those of P microtus are 
more cylindrical, dry with the surface minutely warty and dark brown. 

The villose twigs of Polyalthia lasioclada are similar to those of P. 
borneensis and P. bullata. However these species both have cream to yellow 
flowers with short villose pedicels rather than the orange-red corollas and 
relatively long, sparsely hairy pedicels of P. lasioclada. The many-nerved 
leaves of P. bullata (25-40 pairs) are unlikely to be confused with P. lasioclada 
(15-19 pairs). 


176 Gard. Bull. Singapore 62 (1) 2010 


Acknowledgements 


Many thanks to Dr J.R Veldkamp (L) for translating the Latin diagnosis. 
Support for this reasearch from the Arnold Arboretum, Forest Research 
Institute Malaysia, Singapore Botanic Gardens and Royal Botanic Gardens 
Kew is gratefully acknowledged. 


References 
Johnson, D. M.and N.A. Murray. 1999. Four new species of Polyalthia from 


Borneo and their relationship to Polyalthia insignis. Contributions from 
the University of Michigan Herbarium 22: 95-104. 


Gardens’ Bulletin Singapore 62(1): 177-209. 2010 aT 


Studies on Schismatoglottideae (Araceae) of Borneo XIII: 
A Revision of the Schismatoglottis nervosa 
Species Complex 


WONG SIN YENG 


Department of Plant Science and Environmental Ecology, Faculty of 
Resource Science and Technology, Universiti Malaysia Sarawak, 
94300 Samarahan, Sarawak, Malaysia 


Abstract 


A revision of the Schismatoglottis nervosa Ridl. species complex is presented. 
Ten species are recognized, of which three species are pre-existing (S. 
nervosa, S. elegans A.Hay, and S. brevicuspis Hook.f.) and seven are novel 
and described here (Schismatoglottis adoceta S.Y.Wong, S. linae S.Y.Wong, S. 
matangensis S.Y.Wong, S. simonii S.Y.Wong, S. tessellata S.Y.Wong, S. turbata 
S.Y.Wong, and S. ulusarikeiensis S.Y.Wong.) The S. nervosa species complex 
is readily delimitated by the pungent terpenoid smell when the vegetative 
tissues are crushed, and by the presence of longitudinally ridged petioles. 
A key to the S. nervosa species complex is presented and all species are 
illustrated. 


Introduction 


Schismatoglottis is a genus of in excess of 150 species of terrestrial, lithophytic, 
and rheophytic herbs occurs within Malesia, but mainly in Borneo. Hay & 
Yuzammi (2000) presented an alpha-taxonomic revision of the Malesian 
species, one of the results of which was the recognition of six informal 
species groupings based primarily on shoot architecture and the senescence 
mechanics of the upper spathe: Asperata Group, Calyptrata Group, Corneri 
Group, Multiflora Group, Rupestris Group and Tecturata Group. However, 
Schismatoglottis has been proven as a polyphyletic assemblage (Wong er 
al., 2010), and this has resulted in the removal of the Rupestris Group into 
a resurrected genus, Apoballis (Wong & Boyce, 2010). Further molecular 
analysis is being undertaken and will form the basis of further paper (Ting 
et al., in prep.). 

Hay and Yuzammi (2000) placed S. nervosa and S. elegans (both from 
Sarawak) in the Asperata Group based on the combination of pleionanthic 
shoots, the leaf sheath open and nearly always fully attached and persistent, 
and by the spathe limb opening more-or-less wide and then crumbling- 


178 Gard. Bull. Singapore 62 (1) 2010 


deliquescent. One species, which occurs in West Malaysia, Schismatoglottis 
brevicuspis Hook.f also belongs to this complex. In addition to these 
species, a further 7 species, all novel, are recognized here from Sarawak: 
Schismatoglottis adoceta S.Y.Wong, S. linae S.Y.Wong, S. matangensis 
S.Y.Wong, S. simonii S.Y.Wong, S. tessellata S.Y.Wong, S. turbata S.Y.Wong, 
and S. ulusarikeiensis S.Y.Wong. The S. nervosa species complex is clearly 
distinguished within the Asperata Group by a pungent terpenoid smell 
when crushed (tissues odourless when crushed in Asperata Group s.s¢r., 
i.e., sensu Wong), coriaceous to thinly coriaceous + elliptic leaves with veins 
prominent abaxially (leaves thinly sub-succulent and fragile with veins 
obscure to invisible abaxially in Asperata Group s. str), and longitudinally 
ridged petioles (petioles asperous to puberulent or hispid in Asperata Group 
Ss. Str.), Supporting the recognition as a distinct species complex within the 
group. 

The S. nervosa species complex comprises terrestrial forest-dwelling 
herbs commonly on steep soil banks or occurring as obligate or facultative 
lithophytes on limestones, shales, sandstones and granite. Schismatoglottis 
nervosa and S. elegans are restricted to Karst limestones; Schismatoglottis 
matangensis and S. turbata to sandstones, and S. adoceta, S. tessellata and S. 
ulusarikeiensis to shales, S. simonii to both limestones and sandstones, and S. 
linae and S. brevicuspis to granite. 


Allied taxa 


There are additional species in the S. asperata group that possess 
longitudinally and/or puberulent ridged petioles and/or are weakly (not 
pungently) terpenoid smelling when crushed, of which one, Schismatoglottis 
latevaginata Alderw., described and numerous other species are awaiting 
formal description. Based on initial observations of the spathe senescence 
mechanics (spathe at least partially circumscissile before shedding in large 
pieces) and leaf shape (lamina oblongo-ovate on a disproportionately long 
petiole), these species are morphologically more closely related to one 
another than any of them are to species of the S. nervosa species complex, 
and will be the subject of a subsequent paper. 


Schismatoglottis nervosa species complex 


Small to robust herbs, 40-70 cm tall, occasionally up to ca 1 m tall, with 
vegetative tissues emitting a pungent aromatic (terpenoids) smell when 
crushed. Stem epigeal, pleionanthic, erect to decumbent, sometimes 
ascending and then rooting (‘terrestrial climber’ sensu Boyce), 30-50 cm 


Studies on Schismatoglottideae 179 


long x 2-4 cm thick, adventitiously branched from older portions, rooting 
along entire length on contact with ground, roots often penetrating petiole 
bases; leaf scars prominent. Leaves few to many (5-15) together; innovations 
yellowish green; senescent lamina sometimes rotting and falling together 
with distal portion of petiole to leave the lower portion of the petiole 
attached to plant, this rotting and falling at a later stage; petioles terete, ca 
30-40 cm long, sometimes, up to ca 70 cm long, adaxially channelled or D- 
shaped in cross section, weakly to strongly longitudinally ridged (resembling 
celery - Apium graveolens - Apiaceae) especially abaxially, minutely (strong 
lens required) and densely verruculate or glabrous; petiole sheathing 
in the lower '4- %, sheaths fully attached, thinly coriaceous, sometimes 
leathery, marcescent or persistent, tapering, closed or less often wide open, 
sometimes with a short rounded free ligular portion; lamina broadly ovate 
to oblongo-ovate to elliptic, coriaceous or thinly coriaceous, base broadly 
rounded to sub-truncate, slightly retuse or cuneate, apex acute to strongly 
acuminate, adaxial surface semi-matte, bright deep to medium green, always 
slightly bullate, abaxial surface paler green, often glaucous, drying strongly 
discolorous; midrib adaxially flush with lamina, abaxially very prominent; 
primary venation impressed adaxially, sometimes flush with lamina, strongly 
raised abaxially, alternating with lesser interprimaries, interprimaries 
occasionally arising from the bases of the primary veins, both diverging 
at 45°- 90° and gradually curving towards the apex before reaching the 
intermarginal collecting vein; secondary venation mostly arising from the 
midrib, occasionally from near the bases of the primary veins, prominent 
abaxially; tertiary venation forming an obscure to prominent tessellate 
pattern, variously prominent adaxially and abaxially. Inflorescences up to 
three together (rarely 4), erect, white, moderately fragrant (esterase) at 
female anthesis; peduncle to ca 2 cm long, concealed by leaf bases, prophylls, 
and cataphylls at flowering, slightly exserted in fruit; spathe interior glossy, 
exterior semi-glossy; lower spathe differentiated from the limb by a weak 
constriction correlating with spadix interstice; limb oblongo-lanceolate, white 
to partially green, weakly coriaceous to somewhat spongy, semi-truncate to 
shortly to strongly acuminate (to ca 5 mm long), apex mucronate (to ca 2 
mm), limb either caducous by crumbling at or just after male anthesis, with 
remaining fragments deliquescing, or briefly persistent until end of anthesis 
and thence deliquescent; spadix sessile, isodiametrically adnate on the lower 
abaxial side relative to the spathe opening; female zone conic-cylindric; 
pistils numerous, close-packed or laxly arranged; stigma sessile, punctiform, 
minute to large but always smaller in diameter than ovary; interpistillar 
staminodes confined to a more-or-less single ring at the base of the female 
zone (rarely among the pistils); sterile interstice present, supra-pistillate 
pistillode zone often constricted; pistillodes half to twice the diameter of 


180 Gard. Bull. Singapore 62 (1) 2010 


ovary, close-packed or laxly arranged and followed by a whorl of staminodes, 
this zone often constricted; male zone contiguous with interstice staminodes, 
cylindric; stamens close-packed, the whole butterfly or dumbbell-shaped 
from above; pores oblong to C-shaped, deep to shallow; appendix sub- 
cylindric, white or yellow when fresh, distally tapering to a blunt or sharp 
point, basally merging with the male zone or weakly or distinctly wider and 
slightly truncate; appendix staminodes small to rather large, densely packed, 
flat-topped to slightly impressed, irregularly polygonal. 


Key to Schismatoglottis nervosa species complex 


1. Petiole longitudinally ridged; vegetative tissues pungently aromatic 
(terpenoids) when crushed; leaves + elliptic, thickly to thinly coriaceous 
with vems prominent abaxialll yi ro scscee ss ccsc ae ee eee 2 

1. Not the above combination ......... remainder of genus (see Hay and 
Yuzammi, 2000; Hay, 2002) 


2. Spathe hardly opening; upper part of spathe limb remaining green during 


anthesis; stigma green whemftesins ce:cccse eters eee 5 
2. Spathe opening wide; upper part of spathe turns to white during anthesis; 
stigma white when fresh (except for S. tessellata)) i. secccccess-s-0ee stress - 


3. Petiolar sheath persistent, opening wide; lamina longer and narrower 
(ratio >3:1); primary venation adaxially flush with surface and with less 
than 10-15 on each side. Central Sarawak: Sarikei.......8. ulusarikeiensis 

3. Petiolar sheath marcescent, opening narrow; lamina shorter and wider 
(ratio < 3:1); primary venation adaxially sunken with surface and with 15- 
20 on each side. West Malaysia: widespread.................::00+ S. brevicuspis 


4. Appendix white when fresh; petioles puberulent, densely (sometimes 
minutely) verruculate, longitudinal ridges prominent; leaf apex shortly 
acuminate to Ca’ 1 CIM... <2. ciecccecsectencecepeteenst a eer ee 5 

4. Appendix yellow when fresh; petioles glabrous, not verruculate, 
longitudinal ridges not prominent; leaf apex strongly acuminate to ca 3 


5. Tertiary venation obscure-tessellate; female zone exceeding male 
zone, pistils laxly arranged, stigma small, 1/5 of ovary diam., turning 
yellow in alcohol; spadix interstice sharply constricted. West Sarawak: 
Matai gi siié cc. scssesenseenseosnzenc ene ctess tenet es tetas cestee aes Sess S. matangensis 

5. Tertiary venation prominent tessellate; male zone exceeding female zone, 
pistils crowded, stigma large, 1/3 to % of ovary diam., remaining white 
in alcohol; spadix interstice weakly constricted. Plants of limestone, 
sandstone and granite. West Sarawak except Matang ............c.ceeeeeeees 6 


Studies on Schismatoglottideae 18] 


10. 


Male zone irregularly arranged with uneven surface, male flowers round: 
appendix markedly wider than male zone and slightly truncate basally 
fete MC FET el ATITS OF SATISTOMEs:.-......s005.cccsa+escdescsasscaceecstcstoesenss S. turbata 
Male zone evenly arranged with smooth surface, male flowers butterfly 
shape or dumbbell shape; appendix contiguous with male zone. Plants on 
2) VETTE OLE SUDO IS ICIS Aa a 


Spathe limb splitting into longitudinal strips before deliquescing. Plants 
CL SETILG SIDING soso ee ee S. linae 
Spathe limb deliquescent in a whole piece. Plants of limestone.............. 8 


Stamen pores narrowly C-shaped, deep, connective narrow, rims not 
alate; stigma 2 ovary diam.; secondary and tertiary venation not strongly 
pronounced. Plants weak to robust, sometimes colony forming. Always 
lithophytic in soil collected in crevices and sinkholes in limestone. Full 
shade, less than 200 m asl. West Sarawak: Bau........... eee S. nervosa 
Stamen pores oblong, shallow, connective broad, rims widely alate; stigma 
13 ovary diam.; secondary and tertiary venation pronounced. Plant robust, 
always colony forming. Always terrestrial in deep soil on limestones and 
sandstones. Sometimes in full sun, less than 600 m asl. West Sarawak: 
PVE AC RIR SYST 210 ease so hoe ee S. simonii 


Leaf lamina thinly coriaceous, occasionally with grey stripe, adaxially 
glossy; tertiary venation more tessellate adaxially than abaxially; 
female flowers green; appendix !/ of spadix length. Central Sarawak: 
NGM 222 e doc vont ccc seu sede actbosssvesiSieesonesdcotisnnetenss S. tessellata 
Leaf lamina coriaceous, never variegated, adaxially semi-matte; tertiary 
venation more tessellate abaxially than adaxially; female flowers white; 
ASM ie VARO a Ol SPACES LOMO UN cc cescstseinsdevesstwaceceseesavsosesestacarseecsonentsad 10 


Petiolar sheath leathery; leaf lamina glaucous abaxially, leaf apex shorter 
(to ca 2 cm); stigma large (2 of ovary diameter), staining yellow in alcohol; 
connective narrow, pore oblong. Plants of shales in central and northeast 
SS ll Ad Kept ECS hc Seance de seeen os Eases cose soneoteaceatenseeeshs S. adoceta 


10. Petiolar sheath coriaceous; leaf lamina not glaucous abaxially, leaf apex 


longer (to ca 3 cm); stigma small (1/5 of ovary diameter), remaining white 
in alcohol; connective broad, pore C-shaped. Plants of limestones in 
STINE SE SRE ease Rie eRe ee S. elegans 


1. Schismatoglottis adoceta S.Y.Wong, sp. nov. 

Ab Schismatoglottis elegans foliis subtus glaucis, laminae foliae apice 
acuminata breviore (ad 2 cm longa), stigmatibus maioribus (quam ovaria 
ca 2 diametro vs. 1/5 in S. elegans) et cum alcoholis flavescente (stigmatil S. 
elegansii sum alcoholis alba remenans). Connectivo antherae anguste, poris 


182 Gard. Bull. Singapore 62 (1) 2010 


oblongo (vs. connectivo antherae crasso, poris hippocrepiformis: S. elegans) 
distinguitur. Schismatoglottis adocetae in habitu saxa shaleiaca restrictus est 
(S.elegans in habitu calcicola unica).—Typus: Malaysia, Sarawak, Kapit Division, 
Belaga, km 10 Bakun-Bintulu-Miri road junction, 02° 50° 51.7”; 114° OV 57.6”, 
182 m asl, 11 Oct 2005, PC Boyce et al. AR-1408 (holo, SAR). Plate 1. 


Leaf petioles glabrous, slightly channelled and carinate adaxially in cross 
section; sheathing in the lower \s to 4%, sheath fully attached, closed, tapering, 
leathery, persistent; lamina broadly ovate to oblongo-ovate, more-or-less 
elliptic, thinly coriaceous, slightly bullate, 7-9 cm wide x 13-23 cm long, base 
broadly rounded to sub-truncate, slightly retuse or cuneate, apex acute to 
strongly acuminate to ca 2 cm, adaxial surface semi-matte mid-green, abaxial 
surface paler green, glaucous, drying strongly discolorous, midrib adaxially 
flush with lamina, abaxially prominent, raised canaliculate in cross section; 
primary venation impressed adaxially, strongly raised abaxially with 17-22 
primary lateral veins on each side, alternating with lesser interprimaries, 
frequently with a few branches from near the base, both diverging at 45°- 60° 
and gradually curving towards the apex before reaching the intermarginal 
collecting vein; secondary venation, arising from the midrib and the 
primary veins; tertiary venation obscure tessellate adaxially and abaxially 
in both living plants and dry specimens. Inflorescences up to three in quick 
succession alternating with foliage leaves ca 5 cm long, (1.e. not forming a true 
synflorescence), fragrant (esterase) at female anthesis; peduncle to ca 1 cm, 
not exserted from leaf bases, slightly exserted in fruit; spathe interior glossy, 
exterior semi-glossy, coriaceous, ca 7.5 cm long, lower spathe narrowly ovoid, 
green when fresh, ca 2.5 cm long x | cm diam., differentiated from the limb 
by a weak constriction; limb sub-truncate, white, ca 5 cm long, coriaceous, 
opening wide except for convolute base, at first over-arching, then reflexed, 
finally more or less twisted and then caducous-crumbling, apex mucronate 
ca 4mm long, green when fresh; spadix sessile, adnate isodiametrically to the 
spathe in the lower !/2 of female zone, whitish, to ca 5.5 cm long, less than the 
length of spathe; female zone ca 1.7 cm long, 13 of spadix length; pistils many, 
crowded, round to slightly rhomboid, with weakly bisulcate longitudinal 
ridges, ca 0.5 mm diam. x 0.45 mm long; stigma large, bun-shaped, to ca 0.3 
mm diam., ! of ovary, punctiform, papillate, yellow in alcohol; interpistillar 
staminodes very occasional among the pistils, crowded into a row along 
the spathe/spadix adnation, stalked, clavate and flat-topped, ca 0.5 mm 
diam., similar size to pistils; sterile interstice strongly narrowed, ca 1.4 mm 
diameter in the middle, (2.5-) ca 7 mm long which is 'gof spadix length, 
basally more or less covered with inflated abortive pistils (reduced stigmas), 
twice the diameter of ovary, transversely sulcate, apically covered with two 
rows of sterile stamens, irregular shape, slightly wider than stamens; male 


Studies on Schismatoglottideae 183 


Plate 1. Schismatoglottis adoceta S.Y.Wong. A. Whole plant: B. Leaf lamina adaxially: C. Leaf 
lamina abaxially; D. Female zone; E. Male zone and yellow appendix; F. Inflorescence during 
female anthesis; G. Inflorescence during male anthesis; H. Spadix with the spathe artificially 
removed; I. Spathe at male anthesis; J. Spathe prior to female anthesis. Images © P.C. Boyce 


184 Gard. Bull. Singapore 62 (1) 2010 


zone cylindric, ca 3.6 mm diam. x 1.2 cm long, 13 of spadix length, marked 
increased in diameter from interstice; stamens crowded, truncate, ca 0.4 mm 
across, dumbbell-shaped with the connective not elevated above the thecae, 
connective narrow, pore oblong and deep; appendix sub-cylindric, ca 1.8 
cm long, 1/3 of spadix length, yellow when fresh, turning white in alcohol, 
frequently chewed by beetles, tapering to a sharp point, basally more or less 
not isodiametric with top of male zone; staminodes of appendix more or less 
columnar, irregularly polygonal, flat-topped, ca 0.5-1 mm diameter. 


Other specimens seen: SARAWAK. Bintulu Division: Bintulu, Bk. Satiam, 
02° 59° 33.0”; 112° 56’ 01.4”, 18 m asl, 12 Aug 2004, PC. Boyce & Jeland ak 
Kisai AR-632.1 (SAR); Bk. Merairi, 02° 46’ 26.9”; 113° 39° 19.8”, JS/LC-24 
AR-1284 (SAR). 


Distribution: Central and northeast Sarawak, Kapit and Bintulu Divisions. 
Habitat: Lithophytic on shale (Bintulu and Kapit Divisions), 18-180 m asl. 


Notes: Schismatoglottis adoceta is clearly distinguished from the rest of the 
S. nervosa complex by having leathery petiolar sheath (coriaceous petiolar 
sheath in the rest of the complex). S. adoceta is most similar to S. elegans 
although readily distinguished by the leaf laminae abaxially glaucous, the 
leaf apex much shorter-acuminate (to ca 2 cm), larger stigma (S. adoceta 
12 of ovary diameter vs. S. elegans 1/5 of ovary diameter) staining yellow in 
alcohol (remaining white in S. elegans), anthers with a narrow connective 
and oblong pore (S. adoceta) whereas anthers in S. elegans have a broad 
connective and a C-shaped pore. Schismatoglottis adoceta is restricted to 
shales as compared to S. elegans endemic on limestones. Schismatoglottis 
adoceta differs from S. tessellata (also of shales) by coriaceous leaf laminae 
with the adaxially much less pronounced tertiary venation and the spadix 
appendix in S. adoceta much longer and narrower. 


Etymology: The specific epithet is derived from the Greek ‘adocetus’, 
unexpected, in allusion to the realization late in the preparation of this 
manuscript that several elements of S. elegans sensu Hay and Yuzammi 
required a specific recognition. 


2. Schismatoglottis brevicuspis Hook.f. 

FI. Brit. Ind. 6 (1894) 537; Ridl. Materials Fl. Mal. Pen. 3 (1907) 33 & J. Straits 
Branch Roy. Asiat. Soc. 57 (1910) 113 & Fl. Mal. Pen. 5 (1925) 113; Engl. & 
Krause, Pflanzenr. 55 (I[V.23Da) (1912) 98, pro parte excl. specim. cit. Curtis 
(Penang, Waterfall), Ridley (Selangor, Petaling; Pahang, Tahan River) [i.e. S. 


Studies on Schismatoglottideae 185 


brevipes Hook.f., q.v.]. —Type: Malaysia, Perak, Scortechini 612 (holo, K; iso, 
CAL, SING). 


Other specimens seen: PENINSULAR MALAYSIA. Johor: Muar, Gunung 
Ledang F.R., Gunung Ledang, (Mt Ophir), 22 Jan 1994, (orig. coll. Hay, A., 
Samy, A. & Ban Ka 9172) sub. C. Herscovitch NSW407380 (KEP 41490, L 
0832670); Bukit Tunjok Laut, Ngadiman 37088 (SING). Melaka: Base of G. 
Mering, Ridley s.n. (SING). Pahang: Pulau Tioman, Jason Bay, Burkill 1042 (K, 
SING); Kemaman, Ulu Ayam, Bukit Kajang, Corner 30249e (SING); Raub- 
Bentong boundary, Furtado 33097a (SING). Negeri Sembilan: Beremban 
Forest Reserve, foot of Gunung Angsi, Furtado s.n. (SING). Selangor: Ulu 
Gombak, Croat 53276 (K, MO); Gombak valley, 13 Jan 1994, C. Herscovitch 
NSW407381 (KEP 41493); near Klang Gates Reservoir, Nicolson 1140 (K): 
Genting Peras, Ridley s.n. (SING). Perak: Hulu Perak, Tasik Banding, n.d., 
Baharuddin bin Sulaiman s.n. (sub. AR-2599, living collection in Semenggoh 
Botanical Research, Kuching, Sarawak) (SAR, USM); Sungei Batang Padang, 
Tapah, Furtado 33096 (SING); Larut, 14 Jan 1994, Hay, A., Samy, A. & Ban 
Ka 9075 (L 0832668); Kuala Kangsar, Bubu F.R., foothills of Gunong Bubu, 
18 Jan 1994, (orig. coll. Hay, A., Samy, A. & Ban Ka 9130; Cult. RBG Sydney, 
Acc. No. 940126 (KEP 41475, L 0832671). Kelantan: Gua Musang, Relai FR. 
5°02’ N, 102°23’ E, 1992, Kiew, B.H. KBH10 (KEP 38211). Kedah: Gunung 
Bongsu, nr Terap, Bogner 1692 (K). INDONESIA: Sumatera: Sibolangit, 
Alston 14481 (BM); Sibolangit, Bukit Semiak, Md Nur 7368 (K); Aceh, 
Middle Alas River (Lae Sauraya) area, ca 15 km N of Gelombang, S of 
Bengkong R., 21 Jul 1985, de Wilde & de Wilde-Duyfjes 20188 (L 0239831). 


Distribution: Malay Peninsula and Sumatra (North Sumatera and Aceh 
provinces) 


Habitat: In wet gullies and among rocks by streams in lowland rain forest 
and lowland hill forest, to ca 900 m alt. 


Notes: P.C. Boyce and the author saw a living collection of this species at 
Nancy Botanical Garden, France, and realised that this species belongs to 
the S. nervosa complex, although somewhat unusual in the spathe hardly 
opening and the spathe limb green, a character set otherwise shared only 
with S. uwlusarikiensis. A later trip to the aroid garden in Universiti Sains 
Malaysia, Penang, which also has the living material of S. brevicuspis further 
confirmed the placement of this species. 


3. Schismatoglottis elegans A.Hay 
Telopea 9(1) (2000): 67. Type: Cultivated in RBG Sydney Acc. No. 940510, 


186 Gard. Bull. Singapore 62 (1) 2010 


ex Malaysia, Sarawak, Niah National Park, Niah Caves area, along path 
from Niah town (orig. coll. Hay et al. 9359), Feb 1996, C. Herscovitch s.n. 
(holo, SAR; iso, K, KEP, NSW, all + spirit). Plate 2. 


Plate 2. Schismatoglottis elegans A.Hay. A. Population in the wild, spreading across the floor, 
the base of a limestone hill; B. Whole plant occurs in deep litter on limestone; C. Leaf lamina 
apex (to c.3 cm long); D. Inflorescence at male anthesis; E. Creeping rhizome; F. Inflorescence 
post anthesis showing the deliquescing spathe limb. Images © P.C. Boyce 


Studies on Schismatoglottideae 187 


Other specimens seen: MALAYSIA. Sarawak. Miri Division: Niah Suai, Niah 
National Park, trail to Great Cave, 03° 49’ 09.9”: 113° 46’ 52.3”, 46 m asl, 13 
Oct 2005, P.C. Boyce et al. AR-1428 (SAR); Niah Suai, Niah National Park. 
Madu Trail, 03°48 57.9»; 113° 4618.3», 34 m asl, 13 Jul 2006, PC. Boyce et 
al. AR-1877 (SAR + spirit); P.-C Boyce et al. AR-1878 (SAR + spirit): Subis, 
Gua Sibau, Niah National Park, Trail from Rh. Chang, 24 Aug 2002, Julaihi 
A. et al. S. 89319 (SAR). 


Distribution: Sarawak. Miri Division, endemic to Niah area (Northeast 
Sarawak). 


Habitat: Humus-filled crevices in limestone in wet to swampy lowland 
rainforest, 34-200 m asl. 


Notes: Schismatoglottis elegans is distinguished from S. nervosa (in West 
Sarawak limestone) by leaf laminae not glaucous abaxially and the spadix 
appendix yellow when fresh. The tip of the leaf lamina acuminate up to ca 
3 cm. Stamens of S. elegans are much rounder, with a broad connective and 
are the more dumbbell-shaped as compared to other species in the complex. 
Stamen pores are deepest as compared to other species in the complex. Hay 
and Yuzammi (2000) mentioned that the appendix staminodes dried with 
the stalk collapsed but the tops remaining expanded and tending to cohere 
in groups. However, inflorescence of AR-/877 does not behave in this 
manner while some inflorescences of S. nervosa behave in the same manner 
suggesting that this is not a strong differentiating character. The spadix of 
AR-1877 differs from other collections in that it has ovaries tetrasulcate with 
the stigma staining yellow and interstice staminodes and stamens turning 
darker yellow in alcohol. It is not clear if these are artefacts of the period 
(post anthesis) of preservation. 

Schismatoglottis elegans resembles S. tessellata although aside from 
ecological differences (S. tessellata is restricted to shales) the latter also 
differs by having thinly coriaceous leaf laminae with the tesselate tertiary 
venation adaxially markedly more pronounced, a larger stigma and much 
shorter and broader spadix appendix. 

Hay and Yuzammi (2000) noted that S. elegans is found in a variety 
of other localities in rainforest between 800 and 1300 m alt. However, 
there are considerable doubts about the four specimens cited and pending 
further investigation, the author suspect these observations represent 
morphologically similar but different species. 


4. Schismatoglottis linae S.Y.Wong, sp. nov. 
Schismatoglottis linae simillima speciebus S. nervosa et S. simoni quae 
spathe senescens differt. Spathae laminorum findens longitidinalis ante 


188 Gard. Bull. Singapore 62 (1) 2010 


deliquescens, staminis connectivo producto latus, poris antherae profundis (vs. 
staminis connectivo producto latus, poris non-profundis: S. simonii, et staminis 
connectivo anguste et poris profundis: S. nervosa) distinguitur. Inter specibis 
Schismatoglottodorum grex nervosae habitu graniticola terrestri uniqus. — 
Typus: Malaysia, Sarawak, Kuching Division, Lundu,Gunung Gading, below 
plank walk to swimming area, 01° 41’ 31.0”; 109° 50° 44.5”, 700 m asl, 14 Dec 
2006, P.C. Boyce et al. AR-2062 (holo, SAR). Plate 3. 


Leaf petioles terete, obscurely longitudinally ridged (clearer under 
magnification), glabrous, up to ca 40 cm long, slightly channelled and 
carinate adaxially in cross section; sheathing in the lower 1/3 to %, sheath fully 
attached, closed, tapering, coriouceous, marcescent; lamina broadly ovate to 
oblongo-ovate, more-or-less elliptic, thinly coriaceous, slightly bullate, 8-10 
cm wide x 15-25 cm long, base broadly rounded to sub-truncate, slightly 
retuse or cuneate, apex acute to strongly acuminate to ca 3 cm, adaxial 
surface semi-matte mid-green, abaxial surface paler green, glaucous, drying 
strongly discolorous, midrib adaxially flush with lamina, abaxially prominent, 
raised canaliculate in cross section; primary venation impressed adaxially, 
strongly raised abaxially with 18-22 primary lateral veins on each side, 
alternating with lesser interprimaries, frequently with a few branches from 
near the base, both diverging at 45°- 60° and gradually curving towards the 
apex before reaching the intermarginal collecting vein; secondary venation, 
arising from the midrib and the primary veins; tertiary venation obscure 
tessellate adaxially and abaxially in both living plants and dry specimens. 
Inflorescences up to three in quick succession alternating with foliage leaves 
ca 5 cm long, (i.e. not forming a true synflorescence), fragrant (esterases) at 
female anthesis; peduncle to ca 1 cm, not exserted from leaf bases, slightly 
exserted in fruit; spathe interior glossy, exterior semi-glossy, coriaceous, ca 
13 cm long, lower spathe narrowly ovoid, green when fresh, ca 3.5 cm long 
x 1 cm diam., differentiated from the limb by a constriction; limb truncate, 
white, ca 8 cm long, coriaceous, opening wide except for convolute base, 
at first over-arching, then reflexed, finally more or less twisted and then 
caducous-crumbling in longitudinal stripes, delisquescent post male anthesis, 
apex mucronate ca 4 mm long; spadix sessile, adnate isodiametrically to the 
spathe in the lower 2 of female zone, whitish, to ca 10 cm long, less than the 
length of spathe; female zone ca 2.5 cm long, 1/3 of spadix length; pistils many, 
crowded, round to slightly rhomboid, with weakly bisulcate longitudinal 
ridges, ca 0.5 mm diam. x 0.5 mm long; stigma smaller than ovary, bun- 
shaped, to ca 0.3 mm diam., punctiform, papillate; interpistillar staminodes 
confined into a row along the spathe/spadix adnation, stalked, clavate and 
flat-topped, ca 0.5 mm diam., similar size to pistils; sterile interstice strongly 
narrowed, ca 0.4 cm diameter in the middle, ca 5 mm long, basally more 


Studies on Schismatoglottideae 189 


Plate 3. Schismatoglottis linae S.Y.Wong. A. Whole plant; B. Inflorescence at male anthesis 
with an infructescence on the right side; C. Leaf lamina abaxially; D. Emerging inflorescence 
with two infructescences; E. Inflorescence post anthesis showing the spathe limbs caducous 


in longitudinal stripes. Images © P.C. Boyce 


190 Gard. Bull. Singapore 62 (1) 2010 


or less covered with inflated abortive pistils (reduced stigmas), twice the 
diameter of ovary, transversely sulcate, apically covered with sterile stamens, 
irregular shape, slightly wider than stamens; male zone cylindric, ca 5 mm 
diam. x 3.5 cm long, '4 of spadix length, slightly increased in diameter from 
interstice; stamens crowded, truncate, ca 0.4 mm across, dumbbell-shaped 
with the connective not elevated above the thecae, connective broad, pore 
oblong and deep; appendix subcylindric, ca 3.5 cm long, '4 of spadix length, 
white when fresh and in alcohol, tapering to a sharp point, basally more or 
less not isodiametric with top of male zone; staminodes of appendix more or 
less columnar, irregularly polygonal, flat-topped, ca 0.5-1 mm diameter. 


Distribution: Sarawak, Kuching Division, endemic to Gunung Gading, 
Lundu (West Sarawak). 


Habitat: Terrestrial on pockets of loose soil in deep litter along the stream 
running through at the base of the granites (Kuching Division), 70 m asl. 


Notes: Schismatoglottis linae resembles S. nervosa and S. simonii but is 
distinguished from these two species by the spathe limb senescence. 
The spathe limb splits into longitudinal stripes before deliquescing. The 
connective is broad but stamen pores are deep (broad connective and 
shallow pore in S. simonii, narrow connective and deep pore in S. nervosa). 
Plants are terrestrial along the stream in deep litter on granite surface (the 
only species in the complex on this geology). 


Etymology: This species is named for Lin Jenkins who is an avid lover of 
tropical aroids. 


5. Schismatoglottis matangensis S.Y.Wong, sp. nov. 

Schismatoglottis matangensis cum S. nervosa confunditur sed inflorescentia 
femina quam inforescentia mascula excedens, pistillis sublaxis ordinatis, 
stigmate parva (quam ovaria ca 's diametro) cum alcoholis flavescente, 
spadice cum interstitio sterile abrupte constricta et habitu lapis arenaricola 
restrictus differt. -Typus: Malaysia, Sarawak, Kuching Division, Matang, 
Kubah National Park, Waterfall Trail, 01°35’ 40.2”; 110° 10°45.9”, 190 m asl, 
7 Jun 2006, P.-C. Boyce & S.Y.Wong AR-1830 (holo, SAR). Plate 4. 


Leaf petioles terete, channelled canaliculate adaxially in cross section, 13- 
20 cm long, strongly longitudinally ridged (like celery) especially abaxially, 
very minutely (strong lens required) and densely verruculate, sheathing 
in the lower '4 - % with the sheaths tapering, closed, and fully attached, 
marcescent, sometimes with a short rounded ligular portion; lamina 


Studies on Schismatoglottideae 19] 


Plate 4. Schismatoglottis matangensis S.Y.Wong. A. Whole plant; B. Leaf lamina abaxially; C. 
Inflorescence at male anthesis with an infructescence on the left side; D. Inflorescense at male 
anthesis; E. Female zone, note the stigma in orange; F. Spathe limb deliquescent post male 
anthesis; G. Male zone; H. Spadix with the spathe artificially removed. Images © PC. Boyce 


192 Gard. Bull. Singapore 62 (1) 2010 


broadly ovate to oblongo-ovate, not elliptic, coriaceous, 13-22 cm long x 
7-11 cm wide, the base broadly rounded to slightly retuse, the apex acute 
and shortly acuminate to ca | cm, undulate along margin, adaxial surface 
matte deep green, always slightly bullate, abaxial surface glaucous, paler 
green, drying strongly discolorous; midrib adaxially flush with lamina but 
centrally channelled toward leaf base, abaxially very prominent, raised 
canaliculate in cross section; primary venation impressed adaxially, strongly 
raised abaxially, numerous, 15-22 on each side of the midrib, alternating 
with lesser interprimaries or these occasionally arising from the bases of 
the primary veins, both diverging at 45°-90° and gradually curving towards 
the apex before reaching the intermarginal collecting vein; secondary 
veins mostly arising from the midrib, occasionally from near the bases of 
the primary veins; tertiary venation forming faint tessellate adaxially and 
abaxially. Inflorescences up to 3 together, erect, white, fragrant (esterases) 
smell at female anthesis, peduncle to ca 1.5 cm long, concealed by cataphylls 
deliquesing and adhering to spathe limb exterior, slightly exserted in fruit; 
spathe, interior glossy, exterior, semi-glossy, softly coriaceous, ca 9.5 cm long, 
lower spathe very pale green, ca 4 cm long x 1.3 cm width, differentiated 
from the limb by a faint constriction at interstice, narrowly ovoid; limb 
obovate, white, ca 5.5 cm long x 2.2 cm width, upper margin reflex, abruptly 
acuminate, ca 7 mm long and ultimately mucronate apex, ca 2 mm long, 
caducous by crumbling at or just after male anthesis; spadix adnate, 
isodiametrically attached to 4 of female zone, subcylindric, 5.2 cm long, % 
of spadix length; female zone, conic-cylindric, ivory in fresh and alcohol, 1.6 
cm long x 7 mm diam., '4 of spadix length; pistils many but laxly arranged, 
irregularly round to slight rhomboid from above, ca 0.46 mm diam. x 0.72 
mm long, stigma, sessile, minute, ca 0.15 mm diam., '5 of ovary, punctiform, 
papillate, orange staining in alcohol, ovaries of on lower part zone tending 
to be bisulcate, ovaries of on upper part zone tending to be trisulcate or 
tetrasulcate; interpistillar staminodes confined to less than a single ring at 
the base of the female zone, similar or twice the size of female flower, flat- 
topped: sterile insterstice presents, '/s of spadix length, supra pistilate pistilode 
zone (ca 2.5 mm long), pistilode larger than pistil, irregularly round, 0.6 mm 
diam. transitioning into naked zone (1.5 mm long x 2.3 mm diam.) in AR- 
1830 or contiguous with infra staminate staminodes zone (ca 2.5 mm long), 
irregularly polygonal, flat-topped, white when fresh but stained orange 
in alcohol (AR-1830 & AR-1865), strongly narrow at the staminode zone 
corresponding with the spathe constriction; male zone cylindric, 10 mm long 
x 4mm diam., 4 of spadix length; stamens, butterfly-shaped from above, 
white when fresh, turning dirty yellow in alcohol (AR-1830 & AR-1865), 
remaining white in alcohol (A R-/864), close-packed, 0.3 mm diam., anthers 
truncate (flat-topped), connective narrow and thecae large, pores small and 


Studies on Schismatoglottideae 193 


deep, C-shaped with the convex side innermost; appendix sub-cylindric, ca 
1.9 cm long, *4 of spadix length, white when fresh and in alcohol, strongly 
tapering to a sharp end, middle slightly thicker than the base: staminodes 
of appendix densely packed, flat-topped, irregularly polygonal, ca 0.3-1 mm 
diam. 


Other specimens seen: MALAYSIA. Sarawak. Kuching Division: Matang, 
trail to Indian Temple, 2 Mar 2004, PC. Boyce & Jeland ak Kisai AR-145.3 
(SAR); Matang, Kubah National Park, Waterfall Trail, 01°35’ 40.2”; 110° 
10°45.9”, 190 m asl, 7 Jun 2006, PC. Boyce & S.Y.Wong AR-1831 (SAR); ibid., 
P.C.Boyce & S.Y.Wong AR-1832 (SAR); ibid., PC. Boyce & S.Y.Wong AR- 
1833 (SAR); ibid., PC. Boyce & S.Y¥.Wong AR-1834 (SAR); ibid., 11 Jul 2006, 
P.C. Boyce & S.¥.Wong AR-1864 (SAR + spirit); ibid., PC. Boyce & S.Y.Wong 
AR-1865 (SAR + spirit); ibid., PC. Boyce & S.Y.Wong, AR-1866 (SAR). 


Distribution: Sarawak, Kuching Division, endemic to Matang (West 
Sarawak). 


Habitat: Evergreen moist forest on Matang series sandstones where terrestrial 
in deep litter over exposed black soil, sometimes between sandstone rocks, 
190-450 m asl. 


Notes: Schismatoglottis matangensis is morphologically different from S. 
nervosa by characters of the inflorescence: female zone exceeding male 
zone, pistils laxly arranged [in marked contrast to most other species of the 
S. nervosa complex (except for S. tessellata) in which pistils are crowded] 
and minute (‘4 of ovary diam.) two-three lobed stigma staining yellow 
in alcohol. Other distinguishing characters for S. matangensis include a 
sharp constriction at the interstice, interstice staminodes staining orange 
in alcohol (AR-1830 & AR-1865) and stamens staining orange in alcohol 
(AR-1865) although the remainder of the observed inflorescences have 
stamens remaining white in alcohol and these colour differences may be an 
artefact of inflorescence anthesis phase at preservation although stamens of 
S. nervosa always remain white in alcohol irrespective of the phase of the 
spadix. In robust inflorescences, the appendix displays a marked increase in 
diameter at the junction of the male zone, contiguous with the male but not 
isodiametrical. 

The petiolar sheath is marcescent in S. matangensis. Primary venation 
of S. matangensis is more prominent than in S. nervosa, however, on average 
secondary and tertiary venation are less pronounced. Schismatoglottis 
matangensis is restricted to sandstones whereas S. nervosa are restricted to 
Karst limestones. 


194 Gard. Bull. Singapore 62 (1) 2010 


Etymology: The specific epithet from the Matang Massif, the type locality 
and so far the only known site for this species. 


6. Schismatoglottis nervosa Ridl. 

Schismatoglottis nervosa Ridl., Journ. As. Soc. Straits 49 (1907) 50; Hay 
& Yuzammi, Telopea 9(1) (2000) 73. —Typus: Cult. in Singapore Botanic 
Gardens, ex Malaysia, Sarawak, Bau, Jan 1907, H. N. Ridley s.n. (holo, SING, 
2 sheets). Plate 5. 


Other specimens seen: MALAYSIA. Sarawak. Kuching Division: Bau, Bk. 
Krian, 28 May 1972, J.A.R.Anderson S. 31966 (SAR); Bau, Krokong, Gua 
Peri-peri (Fairy Cave),01° 22° 51.9”; 110° 07? 09:37, 30 measl, 29 Oct 2003; 
PC.Boyce & Jeland ak Kisai AR-145.1 (SAR); Bau, Krokong, Kampung 
Tringgus, Sg. Bong, 01° 157 32.272 110° 05/°37-2-2. 8 aisasle 2 ule O04: 
PC.Boyce & Jeland ak Kisai AR-525 (SAR); Bau, Gn. Bidi, 01° 23’ 27.0”; 
110° 07° 07.6”, 50 m asl, 6 Jan 2005, P.C.Boyce & Jeland ak Kisai AR-944 
(SAR + spirit); Bau, Kampung Bogag, Gn. Tibugai, 01° 21° 31.1”; 110° 03° 
48.7”, 80 m asl, 6 Jan 2005, PC. Boyce & Jeland ak Kisai AR-950 (SAR); Bau, 
Krokong, Gua Peri-Peri (Fairy Cave), 01° 22’ 51.9”; 110° 07’ 09.3”, 30 m asl, 
25 May 2006, P.C.Boyce & S.Y.Wong AR-1823 (SAR); ibid., P-C.Boyce & 
S.Y¥. Wong AR-1824 (SAR); Ibid., RC. Boyce & S.Y.Wong AR-1825 (SAR); 
ibid., PC.Boyce & S.Y.Wong AR-1826 (SAR + spirit); Ibid., PC. Boyce & 
S.Y¥.Wong AR-1827 (SAR + spirit); Bau, Kampung Jugan, 01° 28° 46.4”; 110° 
0S’ 08.5”, 72 m asl, 25 Jun 2004, P.-C. Boyce et al. AR-491 (SAR); Bau, Gn. 
Singai, 17 Nov 2004, PC. Boyce etal. AR-752 (SAR); Bau, Krokong, Kampung 
Tringgus, 01° 15’ 40.2”; 110° 05’ 35.9”, 80 m asl, 19 Feb 2005, PC Boyce et al. 
AR-992 (SAR); Bau, Gn. Juita, 01° 23’ 48.7”; 110° 08’ 07.2”, 35 m asl, 28 
Oct 2005, P.-C. Boyce et al. AR-1499 (SAR); Bau, Segong, Gn. Opar, 01° 27° 
07.3”; 110° 04’ 00.5”, 79 m asl, 9 Nov 2005, P.-C. Boyce et al. AR-1502 (SAR); 
Bau, Krokong, Kampung Tringgus, 01° 15’ 40.2”; 110° 05° 35.9”, 81 m asl, 27 
Jun 2006, P.C.Boyce et al. AR-1845 (SAR + spirit); Bau, Gn. Lanyang, 10 
Apr 2002, Connie, G et al. SBC 2807 (SAR); Bau, Kampung Jugan, 19 Jun 
2004, Jeland ak Kisai & Jipom ak Tisai AR-474 (SAR); Bau, Gn. Poing, 23 
Sep 2001, Julia, S. et al. SBC 345 (SAR); locality not recorded, 26 Feb 2002, 
Julia, S. et al. SBC 2155 (SAR); Bau, Bk. Jebong, 6 Jul 1970, RE Lehmann 
S. 30136 (SAR); Bau, Jambusan, Gn. Batu, 19 Feb 2002, K. Meekiong et al. 
SBC 1662 (SAR); Bau, Jambusan, Gn. Batu, 19 Feb 2002, K. Meekiong et al. 
SBC 1929 (SAR); Bau, Jambusan, Gn. Jebong, 5 Mar 2002, K. Meekiong et 
al. SBC 1953 (SAR); Bau, Bengoh range, Pangkalan Tebang, logging road, 
6 Jul 1996, M. Mohizah S 73890 (SAR); 2 miles east of Bau, 6 Aug 1961, D. 
H. Nicolson (SAR, US); Bau, Krokong, Gua Peri-Peri (Fairy Cave), 22 Mar 
1999, Patsipun et al. S. 79985 (SAR); Bau, Gn. Tabai, 13 Mar 2002, Shaevy W. 
et al. SBC 2298 (SAR). 


Studies on Schismatoglottideae 195 


Plate 5. Schismatoglottis nervosa Ridl. A. Whole plant; B. Inflorescence prior to anthesis: 
C. Inflorescence at male anthesis: D. Whole plant with an inflorescence at male anthesis: E. 
Spadix with the spathe artificially removed; F. Female zone: G. Spathe limb deliquescent post 
anthesis. Images © P.C. Boyce 


196 Gard. Bull. Singapore 62 (1) 2010 


Distribution: Sarawak, Kuching Division, endemic to the Bau area (West 
Sarawak). 


Habitat: Evergreen moist forest on karst limestone in the Bau area. Terrestrial 
in deep litter or lithophytic in soil collected in crevices and sinkholes in 
limestone; rarely along stream banks, 30-160 m asl. 


Notes: Hay and Yuzammi (2000) were unclear whether the interstice 
was present or the fertile zones were contiguous. Based on the author’s 
observation, the interstice is always present with pistillodes at the lower 
part and staminodes at the upper part and without the presence of naked 
zone. In the type specimen, the presence of the interstice is not clear due to 
damage to the spadix at the crucial area of the interstice. In all collections 
the female zone is shorter than the male zone, whereas S. matangensis has 
a longer female zone than male zone. Stigmas of S. nervosa are larger than 
those of S. matangensis and always remain white in alcohol. 

In the type specimen, there are two types of staminodes on the 
appendix of the inflorescence. The lower staminodes are very irregularly 
rhomboid, smaller (ca 0.5 mm diam.) while the upper staminodes are 
regularly rhomboid and larger (0.5-1 mm diam.). A naked zone, which has 
dried black, is present at the tip of appendix. 


7. Schismatoglottis simonii S.Y.Wong, sp. nov. 

A Schismatoglottis nervosa habitu robustis coloniis, veneris laminorum 
seconadariis et tertiariis plus prominentis, stigmate plus late, connectivo 
antherae crasso, poris oblongo, non profundus quam marginem late alatis 
differt. -Typus: Malaysia, Sarawak, Samarahan, Serian, Kidadum, Sugun 
Karang, 01°06’ 17.6”; 110° 29°04.5”, 100 m asl, 29 Jun 2006, P.C Boyce et al. 
AR-1859 (holo, SAR). Plate 6. 


Leaf petioles terete, channelled slightly canaliculate adaxially in cross 
section, 13-40 cm long, strongly longitudinally ridged (like celery) especially 
abaxially at the base, very minutely (strong lens required) and densely 
verruculate, sheathing in the lower '4- % with the wings tapering, and fully 
attached, marcescent,sometimes with a short rounded ligular portion; lamina 
broadly ovate to oblongo-ovate, sometimes elliptic, 12-30 cm long x 6-20 
cm wide, coriaceous, adaxial surface semi-glossy deep green, always slightly 
bullate, abaxial surface paler green, glaucous, drying strongly discolourous, 
the base broadly rounded to sub-truncate, slightly retuse to cuneate, the 
apex acute and shortly acuminate to ca | cm, undulate along margin; midrib 


= 4 : a 
Studies on Schismatoglottideae 197 


Plate 6. Schismatoglottis simonii S.Y.Wong. A. Whole plant; B. Inflorescence during male 
anthesis; C. Leaf lamina adaxially; D. Leaf lamina abaxially: E. Emerging inflorescence with 
three infructescense flanking on each side; F. Spadix with spathe artificially removed: G. 
Female zone; H. Spathe limb starting to deliquescent post male anthesis; I. Infructescence 
ripen to reveal the seeds. 


198 Gard. Bull. Singapore 62 (1) 2010 
ip 


adaxially flush with the lamina but centrally channelled toward the leaf base, 
abaxially very prominent raised canaliculate; primary venation impressed 
adaxially, strongly raised abaxially, numerous, 16-25 on each side of the 
midrib, alternating with lesser interprimaries or these occasionally arising 
from the bases of the primary veins, diverging at 60°-90° and gradually 
curving towards the tip before reaching the intermarginal colllecting vein; 
secondary veins mostly arising from the midrib, occasionally from near the 
bases of the primary veins; tertiary venation forming tessellate reticulum 
adaxially and abaxially, more notable abaxially. Inflorescences up to 
three together, erect, white, fragrant (esterases) smell at female anthesis; 
subtended by lanceolate cataphylls to ca 6 cm long; peduncle to ca 2.5 cm 
long, concealed by leaf bases and cataphylls at flowering, slightly exserted in 
fruit; spathe, interior glossy, exterior semi-glossy, softly coriaceous, ca 9 cm 
long; lower spathe, ovoid, pale green when fresh and white in alcohol, ca 2.5 
cm long x 1.5 cm diam., differentiated from the limb by a faint constriction 
coinciding with interstice; limb oblongo-lanceolate, white when fresh and 
in alcohol, ca 5.5 cm long, apex, green when fresh, mucronate for ca 5.8 
mm, caducous by disintegrating or crumbling at or just after male anthesis; 
spadix sessile, adnate isodiametrically to the spathe in the lower '/ of female 
zone, subcylindric, ca 7 cm long subequalling the spathe; female zone ovoid, 
ivory when fresh and in alcohol, 1.8 cm long x 0.8 cm diam., '4 of spadix 
length; pistils numerous and close-packed, rhomboid to round from above, 
ca 1.19 mm long x 0.56 mm diam.; stigma sessile, punctiform, large, 0.20 mm 
diam., '4 of ovary, raised; interpistillar staminodes confined to more or less a 
single ring at the base of the female zone, irregularly polygonal from above, 
about the same size as the ovaries shorter than pistils; sterile insterstice 
present, supra pistillate pistillodes zone, white, ca 1.8 mm long, '/4 of spadix 
length, strongly narrow in the upper half of pistillodes zone corresponding 
with the spathe constriction, pistillodes irregularly polygonal, large, ca 1 mm 
diam., twice the size of ovary, centrally impressed; sometimes naked zone 
present; followed by a whorl of incompletely abortive stamens, ca 2.6 mm 
long, staminodes, irregularly squat-columnar, flat-topped, 0.62 mm diam.; 
male zone contiguous with interstice staminodes, cylindric, 1.3 cm long x 
4.2 mm diam., slightly less than '4 of spadix length; stamens close-packed, 
irregular dumbbell-shaped from above and neighbouring anthers with their 
lobes interdigitating, white when fresh and remaining white in alcohol, ca 
0.6 mm diam., anthers truncate, connective broad and elevated, thecae large; 
pores oblong and shallow, rims widely alate; appendix cylindric, 2.6 cm long 
x 4.8 mm diam., slightly less than '4 of spadix length, white when fresh and in 
alcohol, abruptly widen at base, distally tapering to sharp point; staminodes 
of appendix densely packed, flat-topped to slightly centrally impressed, 
irregularly polygonal, ca 0.5-1 mm diam. 


Studies on Schismatoglottideae 199 


Other specimens seen: MALAYSIA. Sarawak, Kuching Division: Padawan, 
Kampung Bengoh, Danu road, Gn. Temuang, Sg. Abang, 01° 15’ 38.6”; 110° 
15° 31.4”, 50 m asl, 16 Feb 2006, PC. Boyce et al. AR-1707 (SAR); Padawan, 
10 mi s.w. of main Kuching-Serian Highway, 01° 10°; 110° 20°, 30 Sep 1981, 
T: B.Croat 53179 (SAR); Siburan, Kampung Giam, Air Terjun Giam, 01° 19” 
11.2”; 110° 16° 11.4”, 37 m asl, 7 Feb 2006, P.-C. Boyce et al. AR-1693 (SAR); 
Samarahan Division: Serian, Mongkos, Kampung Batuh Mawang, Labak 
Ebang, Utak Samat, 5 Jan 2006, Simon Kutuh ak Paru AR-1666 (SAR); 
Serian, Gn. Ampungan, 01° 09’ 08.2”; 110° 37° 21.2”, 450 m asl, 21 Nov 2003, 
P.C.Boyce & Jeland ak Kisai AR-92.4 (SAR); Serian, Pichin, Umon Murut, 
Tiab Belanting, 01° 08’ 03.7”; 110° 27’ 00.3”, 90 m asl, 15 Jun 2005, P.C. Boyce 
et al. AR-1215 (SAR); ibid., 15 Jun 2005, P.-C. Boyce et al. AR-1220 (SAR + 
spirit); Serian, Mongkos, Kampung Batuh, Gn. Selabur, 00°57° 26.2”; 110° 
30’ 15.8”, 100 m asl, 15 Mar 2006, PC. Boyce et al. AR-1724 (SAR); Serian, 
Kidadum, Sugun Karang, 01°06’ 17.6”; 110° 29°04.5”, 100 m asl, 7 Apr 2006, 
P.C.Boyee et al. AR-1764 (SAR); Serian, Gn. Ampungan, 01° 09’ 10.1”; 110° 
37 26.2”, 568 m asl, 28 Aug 2006, P.-C. Boyce et al. AR-2003 (SAR + spirit); 
Serian, Pichin, Gunung Kedadum, Sugun Kerang, 13 Nov 2004, Simon Kutuh 
ak Paru AR-750 (SAR); Serian, Pichin, Labu, Sg. Tiyab, 26 Jul 2005, Simon 
Kutuh ak Paru AR-1299 (SAR); Serian, Kampung Selabi, Sg. Mawang, 2 
Feb 2006, Simon Kutuh ak Paru AR-1703 (SAR): Serian, Taman Rekreasi 
Rachan, 01° 08’ 34.9”; 110° 35’ 02.4”, 57 m asl, 18 Oct 2006, PC. Boyce & 
S.Y.Wong AR-2038 (SAR). 


Distribution: Sarawak, Kuching & Samarahan Divisions, endemic to the 
Padawan/Serian areas (West Sarawak). 


Habitat: Always terrestrial in deep soil on limestones and sandstones, 
sometimes lithophytic on permanent wet areas of limestone but not in 
direct water flow, sometimes not in full shade. Large colony forming, 50-600 
m asl. 


Notes: Schismatoglottis simonii is clearly closely allied to S. nervosa but leat 
laminae are wider in S. simonii than S. nervosa [{length: width ratio (to 3:2 
in S. simonii, to 2:1 in S. nervosa)]. Secondary and tertiary venations in S. 
simonii are more pronounced than S. nervosa. The zonation in the spadix 
of S. simonii is equally distributed among the female, male and appendix 
(‘seach) as compared to S. nervosa (female zone, 4; male zone *s; appendix 
',). The stigma narrower in S. simonii as compared to S. nervosa while the 
stamens have a broad connective, with shallow oblong pores. S. simonii tends 
to form large colony and much more robust than S. nervosa. 


200 Gard. Bull. Singapore 62 (1) 2010 


Etymology: This species is named for Mr Simon Kutuh ak Paru, occasional 
member of our field team, who has so ably organise field trips to Padawan 
limestones. 


8. Schismatoglottis tessellata S.Y.Wong, sp. nov. 

Folit lamina tenuiter coriaceis, nervis tertiariis tessellatis abaxialiter et 
adaxialiter valde prominentibus quamquam veniis adaxialiter valde 
prominentibus, stigmate amplis globuliformis (quam ovaria ca 42 diametro); 
appendice crasso brevioribus, usque ad tertiam partem longitudinis spadice 
toto in complexus nervosae unica est. In habitu saxa shaleiaca restrictus. — 
Typus: Malaysia, Sarawak, Kapit, Taman Rekreasi Sebabai, 01° 56° 45.6”; 
112° 54 16.8”, 50 m asl, 16 Mar 2005, P.-C. Boyce et al. AR-1087 (holo, SAR). 
Plate 7. 


Leaf petioles, slightly D-shaped but slightly carinate channelled to the 
base of lamina adaxially in cross section, glabrous, longitudinal ridges only 
visible with magnification, long, 20-30 cm long; sheathing in the lower ‘4 to 
¥Y2 with the sheaths tapering, fully attached, open, persistent to marcescent; 
lamina broadly ovate to oblong ovate, thinly coriaceous, 13-21 cm long x 5- 
10 cm wide, sometimes variegated with grey stripes on each side adaxially, 
the base cuneate to slightly retuse, the apex acute and strongly acuminate 
for ca 0.7-2 cm, adaxial surface semi-glossy deep green, abaxial surface 
paler green, always slightly bullate, glaucous, drying strongly discolourous; 
midrib adaxially flush with the lamina but centrally channelled at the leaf 
base, abaxially very prominent, raised canaliculate in cross section; primary 
venation impressed adaxially, strongly raised abaxially, numerous, 11-20 
on each side of the midrib, alternating with lesser interprimaries or these 
occasionally arising from the bases of the primary veins, diverging at 45°- 
60° and gradually curving towards the tip before reaching the intermarginal 
collecting vein; secondary veins mostly arising from the midrib, occasionally 
from near the bases of the primary veins; tertiary venation forming distinctive 
tessellate adaxially and abaxially, more notable adaxially. Inflorescences up 
to three together, erect, white, fragrant (esterases) smell at female anthesis; 
concealed by oblongo-lanceolate cataphyll,ca 7 cm long, peduncle to ca 2 cm, 
slightly exserted in fruit; spathe, interior glossy, exterior semi-glossy, softly 
coriaceous, ca 9 cm long; lower spathe broadly ovoid, ca 1.7 cm diam. x 3.8 
cm long, differentiated from the limb by a faint constriction coinciding with 
interstice; limb broadly lanceolate, rather coriaceous, caducous by crumbling 
at or just after male anthesis; spadix, sessile, isodiametrically attached to the 
spathe in the lower '4of female zone, conic-cylindric, ca 5.9 cm long; female 
zone, conic-cylindric, white when fresh but very slightly yellow in alcohol, 1.4 


Studies on Schismatoglottideae 201 


Plate 7. Schismatoglottis tessellata S.Y.Wong. A. Whole plant on shales; B. Leaf lamina 
adaxially revealing the tessellate venation; C. Leaf lamina variation; D. Inflorescence at male 
anthesis; E. Spathe at male anthesis; F. Spadix with spathe artificially removed; G. Female 
zone, interstice and part of male zone. Images © P.C. Boyce 


202 Gard. Bull. Singapore 62 (1) 2010 


cm long x 7 mm diam., 4% of spadix length; pistils numerous, laxly-arranged, 
elongate-ovoid to subcylindric, ca 1.70 mm long x 0.53 mm diam.; stigma 
sessile, large bun-shaped, ca 0.26 mm diam., 2 of ovary, raised, papillate; 
interpistillar staminodes confined to more or less a single ring at the base of 
the female zone, irregularly polygonal from above, twice the size of ovaries, 
flat-topped, shorter than pistils; sterile interstice presence, 1.2 cm long, '/ 
of spadix length, pistils transitioning gradually into laxly spirally arranged 
pistilode that are ca half to two times larger than pistil, irregularly round, 
white when fresh and in alcohol, ca 7 mm long, sharp constriction in between 
pistillode and staminode zones; staminodes, larger than stamens, irregular 
polygonal, flat-topped, white when fresh and turning yellow in alcohol, ca 
5 mm long; male zone, cylindric, ca 2.2 cm long x 4.8 mm diam., '4 of spadix 
length; stamens close-packed, irregularly butterfly-shaped from above and 
neighbouring anthers with their lobes interdigitating, 0.5 mm diam., anthers 
truncate, connective broad and thecae large, pores large accounting of the 
entire top of the thecae and shallow, C-shaped, the rims narrowly alate; 
appendix conic-cylindric, 1.2 cm long x 5.3 mm diam., 4 of spadix length, 
yellow when fresh, turning white in alcohol, distally tapering to a very blunt 
point, basally slightly thicker than the male zone, middle thicker than basal; 
staminodes of appendix loosely arranged, large staminode, flat-topped, 0.7- 
1.2 mm across, squat-columnar. 


Other specimens seen: MALAYSIA. Sarawak, Kapit Division: Nanga Gaat, 
Rejang Wood Concession, Sg. Piat, 01° 38’ 09.1”; 113° 24’ 09.9”, 200 m asl, 
14 Oct 2003, P.C.Boyce & Jeland ak Kisai AR-103.1; P.-C. Boyce & Jeland ak 
Kisai AR 103.2 (SAR); ibid., 14 Oct 2003, P.C. Boyce & Jeland ak Kisai AR- 
105 (SAR); Nanga Gaat, km 3.5 after heli-logging camp on road to Camp 
Gahada, Sg. Bereng, 01° 45’ 36.0”; 113° 27’ 54.7”, 228 m asl, 15 Dec 2004, 
P.C. Boyce et al. AR-888 (SAR); ibid., 19 Apr 2006, P.-C. Boyce et al. AR-1792 
(SAR); ibid., 19 Apr 2006, P.-C. Boyce et al. AR-1794 (SAR). 


Distribution: Central Sarawak (Kapit Division). 


Habitat: Evergreen moist forest on shale where either on steep earth banks 
or lithophytic on muddy shale, 50-228 m asl. 


Notes: This species is distinguishable by its thinly coriaceous leaves texture 
and tertiary venation strongly raised-tessellate in both surfaces of leaf, 
but more prominently adaxially. The pistils are laxly arranged with large 
(compared to ovary diameter) bun-shaped stigmas, while the appendix is 
shorter and broader ( ' of spadix length) than other species of the S. nervosa 
species complex. Schismatoglottis tessellata bears some resemblance to S. 


Studies on Schismatoglottideae 203 


elegans but is readily separable by its thinly coriaceous leaf lamina, adaxially 
strongly prominent tessellate tertiary venation (hence the epithet) and 
shorter leaf apex. The stigma is large bun-shaped in S. tessellata as compared 
to small stigma in S. elegans. Schismatoglottis tessellata is restricted to shales, 
as observed so far. Occasionally, leaf laminae of S. tessellata are variegated 
with grey stripes on each side adaxially. 


Etymology: The epithet reflects the prominently raised-tessellate venation 
that immediately distinguishes this species. 


9. Schismatoglottis turbata S.Y.Wong, sp. nov. 

Schismatoglottis turbata ad alii specibis Schismatoglottodorum grex floribus 
masculinus parvus, rotundis, densiter irregularis dispositae, poris profunde 
differt. Appendice quam inflorescentiae masculae apicem sigillatim crassiore 
distinguitur. —Typus: Malaysia, Sarawak, Kuching Division, Sempadi, Sg. 
Limau, Bukit Kankar, 01°39" 44.2”; 109°59°56.5”, 41 m asl, 25 Aug 2007, 
P.-C. Bovyee et al. AR-2143 (holo, SAR). Plate 8. 


Leaf petioles terete, obscurely longitudinally ridged (clearer under 
magnification), glabrous, up to ca 30 cm long, slightly channelled and 
carinate adaxially in cross section; sheathing in the lower ‘4 to 4, sheath 
fully attached, closed, tapering, leathery, persistent; lamina broadly ovate 
to oblongo-ovate, more-or-less elliptic, thinly coriaceous, slightly bullate, 7- 
9 cm wide x 13-23 cm long, base broadly rounded to sub-truncate, slightly 
retuse or cuneate, apex acute to strongly acuminate to ca 2 cm, adaxial 
surface semi-matte mid-green, abaxial surface paler green, glaucous, drying 
strongly discolorous, midrib adaxially flush with lamina, abaxially prominent, 
raised canaliculate in cross section; primary venation impressed adaxially, 
strongly raised abaxially with up to 20 primary lateral veins on each side, 
alternating with lesser interprimaries, frequently with a few branches from 
near the base, both diverging at 45°- 60° and gradually curving towards the 
apex before reaching the inter-marginal collecting vein; secondary venation, 
arising from the midrib and the primary veins; tertiary venation obscure 
tessellate adaxially and abaxially in both living plants and dry specimens. 
Inflorescences up to four in quick succession alternating with foliage leaves 
ca 5 cm long, (i.e. not forming a true synflorescence), fragrant (esterases) at 
female anthesis; peduncle to ca 1 cm, not exserted from leaf bases, slightly 
exserted in fruit; spathe interior glossy, exterior semi-glossy, coriaceous, ca 
7.5 cm long, lower spathe narrowly ovoid, green when fresh, ca 2.5 cm long 
x 1 cm diam., differentiated from the limb by a weak constriction: limb sub- 
truncate, white, ca 5 cm long, coriaceous, opening wide except for convolute 
base, at first over-arching, then reflexed, finally more or less twisted and 


204 Gard. Bull. Singapore 62 (1) 2010 


Plate 8. Schismatoglottis turbata S.Y.Wong. A. Whole plant; B. Two emerging inflorescences 
with two infructescences; C. Inflorescence at male anthesis; D. Spadix with spathe artificially 
removed; E. Female flowers, interstice and part of male zone; F. Male zone and appendix; G. 
Spathe limb deliquescent post anthesis. Images © P.C. Boyce 


Studies on Schismatoglottideae 205 


then caducous-crumbling, apex mucronate ca 4 mm long, green when 
fresh; spadix sessile, adnate isodiametrically to the spathe in the lower ‘4 
of female zone, whitish, to ca 5.5 cm long, less than the length of spathe: 
female zone ca 1.7 cm long, 4 of spadix length; pistils many, crowded, round 
to slightly rhomboid, with weakly bisulcate longitudinal ridges, ca 0.5 mm 
diam. x 0.45 mm long; stigma large, bun-shaped, to ca 0.3 mm diam., ‘sof 
ovary, punctiform, papillate, yellow in alcohol; interpistillar staminodes very 
occasional among the pistils, crowded into a row along the spathe/spadix 
adnation, stalked, clavate and flat-topped, ca 0.5 mm diam., similar size to 
pistils; sterile interstice strongly narrowed, ca 1.4 mm diameter in the middle, 
(2.5-) ca 7 mm long which is ‘4 of spadix length, basally more or less covered 
with inflated abortive pistils (reduced stigmas), twice the diameter of ovary, 
transversely sulcate, apically covered with two rows of sterile stamens, 
irregular shape, slightly wider than stamens; male zone cylindric, ca 3.6 mm 
diam. x 1.2 cm long, '4 of spadix length, marked increased in diameter from 
interstice; stamens crowded, truncate, ca 0.4 mm across, dumbbell-shaped 
with the connective not elevated above the thecae, connective narrow, pore 
oblong and deep; appendix subcylindric, ca 1.8 cm long, '4 of spadix length, 
yellow when fresh, turning white in alcohol, frequently chewed by beetles, 
tapering to a sharp point, basally more or less not isodiametric with top 
of male zone; staminodes of appendix more or less columnar, irregularly 
polygonal, flat-topped, ca 0.5-1 mm diameter. 


Other specimens seen: MALAYSIA. Sarawak, Kuching Division, Sempadi, 
Sg. Limau, 250 m asl, 26 Mar 2004, RC. Boyce & Jeland ak Kisai AR-271 
(SAR). 


Distribution: Southern Sarawak, Kuching Division. 
Habitat: Lithophytic on sandstone (Kuching Division), 41-250 m asl. 


Notes: Schismatoglottis turbata is clearly distinguished from the rest of the S. 
nervosa complex by having small and round male flowers with deep pores. 
The male zone is dense but irregularly arranged. The appendix is markedly 
increased in diameter than the male zone. 


Etymology: The specific epithet is derived from the Latin ‘twrbatus’, 
exasperating, in allusion to the feeling on discovering yet another novel 
species in the S. nervosa complex. 


10. Schismatoglottis ulusarikeiensis S.Y.Wong, sp. nov. 
Ab alii speciebus grex nervosae Borneensibus spathae laminorum pro parte 


206 Gard. Bull. Singapore 62 (1) 2010 


majore viridis (non in toto albis) et per anthesin haud aperiens distinguitur. 
~Typus: Malaysia, Sarawak, Sarikei Division, Ulu Sarikei, 01° 55’ 05.4”; 111° 
29’ 35.8”, 59 m asl, 7 Dec 2005, P.-C. Boyce et al. AR-1588 (holo, SAR). Plate 9. 


Leaves few together (to ca 5), innovations yellowish green, the lamina 
sometimes rotting and leaving behind portion of petiole, with this rotting 
away at a later stage; petioles terete, channelled slightly canaliculate adaxially 
in cross section, sometimes D-shaped in cross section, strongly longitudinally 
ridged, very minutely (strong lens required) and densely verruculate, 13-20 cm 
long, sheathing in the lower '/3 to % with the sheaths tapering, fully attached, 
wide open persistent; lamina always oblongo-ovate, sometimes ovate, softly 
coriaceous, 13-21 cm long x 5-9 cm wide (ratio up to ca 3:1), the base broadly 
rounded to sub-truncate, slightly retuse to cuneate, the apex acute to ca 1 
cm, adaxial surface semi-glossy mid green, abaxial surface paler green, always 
slightly bullate, glaucous, drying strongly discolourous; midrib adaxially 
flush with the lamina but centrally channelled at the leaf base, abaxially very 
prominent, raised canaliculate in cross section; primary venation prominent 
but flushed on adaxial surface, sometimes slightly impressed, strongly raised 
abaxially, numerous, 10-15 on each side of the midrib, alternating with lesser 
interprimaries or these occasionally arising from the bases of the primary 
veins but only occur at the leaf bases, diverging at 60°- 80° and gradually 
curving towards the tip before reaching the intermarginal collecting vein; 
secondary veins mostly arising from the midrib, occasionally from near the 
bases of the primary veins, obscure adaxially and abaxially; tessellate tertiary 
venation, obscure adaxially and prominent abaxially. Inflorescences up to 
three together, erect, white, 6.5 to 7 cm long, peduncle 1.5 to 2.5 cm, slightly 
exserted during fruiting; spathe, interior glossy, exterior semi-glossy, thickly 
coriaceous, ca 9 cm long; lower spathe broadly ovoid, ca 1.5cm diam. x 3.5 cm 
long, white when fresh, differentiated from the limb by a faint constriction 
coinciding with the upper part of interstice; limb lanceolate, thickly coriaceous, 
caducous by melting at or just after male anthesis, upper part of the limb 
green prior to anthesis; spadix, sessile, isodiametrically attached to the spathe 
in the lower '/s of female zone, cylindric, 5.5-6 cm long; female zone, cylindric, 
green when fresh but yellow in alcohol, 1.5-2.2 cm long x 7-10 mm diam., 
’ of spadix length; pistils numerous, tightly arranged, elongate-ovoid to 
subcylindric, ca 0.50 mm diam.; stigma sessile, large bun-shaped, ca 0.40 mm 
diam., overtopping ovary, raised, papillate; interpistillar staminodes confined 
to more or less a single ring at the base of the female zone, but sometimes in 
robust inflorescence, a few scattering towards the distal part of the female 
tzone, irregularly polygonal from above, twice the size of ovaries, flat-topped, 
shorter than pistils; sterile interstice presence, 0.6-1 cm long x 0.7-1 cm wide, 


Studies on Schismatoglottideae 207 


Plate 9. Schismatoglottis ulusarikeiensis S.Y.Wong. A. Whole plant: B. Petiolar sheath 
persistent and open; C. Inflorescence at just before female anthesis; D. Female zone, interstice 
and part of male zone: E. Spathe prior to anthesis; F. Spadix with spathe artificially removed: 
G. Inflorescence post male anthesis with the appendix and spathe limb deliquescent. 


208 Gard. Bull. Singapore 62 (1) 2010 


pistillodes that are c. half to two times larger than pistil, irregularly round, 
white when fresh and in alcohol, ca 7 mm long, sharp constriction in between 
pistillode and staminode zones; staminodes, larger than stamens, irregular 
polygonal, flat-topped, white when fresh and turning yellow in alcohol, ca 5 
mm long; male zone, cylindric, ca 1.8-2.2 cm long x 7 mm diam., '4 of spadix 
length; stamens close-packed, irregularly butterfly-shaped from above and 
neighbouring anthers with their lobes interdigitating, 0.5 mm diam., anthers 
truncate, connective narrow and thecae large, pores large accounting of 
the entire top of the thecae and shallow, C-shaped, the rims narrowly alate; 
appendix cylindric, 2.1-2.3 cm long x 7 mm diam., '4 of spadix length, white 
when fresh and in alcohol, blunt point at distall, basally contiguous with the 
male zone; staminodes of appendix tightly arranged, large staminode, flat- 
topped, 0.7-1.2 mm across, squat-columnar, staminodes at the distal portion, 
sometimes forms slit. Fruits immature during observation. 


Other specimens seen: MALAYSIA. Sarawak, Sarikei Division: Ulu Sarike1;01° 
55’ 05.4”; 111° 29” 35.8”; 59 m asl, 7 Dec 2005, P.-C. Boyce et al. AR-1577 (SAR); 
ibid., 7 Dec 2005, R-C. Boyce et al. AR-1579 (SAR); ibid., 7 Dec 2005, PC. Boyce 
et al. AR-1588 (SAR) ; ibid., 7 Dec 2005, PC. Boyce et al. AR-1635 (SAR). 


Distribution: Central Sarawak, Sarikei Division. Known only from the type 
locality. 


Habitat: Evergreen moist lowland forest on shales, 59 m asl. 


Notes: Schismatoglottis ulusarikeiensis is immediately distinguishable from 
the rest of the S. nervosa complex with primary venation adaxially flush with 
surface and with less than 10-15 on each side. Petioles are terete, strongly 
longitudinally ridged and without hyaline margin in S. ulusarikeiensis. The 
lamina is longer but narrower as compared to the rest of the species in the 
complex. The upper part of the spathe limb remains in green form prior to 
anthesis and this is unobserved in any other species in the complex. The 
spathe limb is barely open during anthesis and this is only observed in S. 
brevicuspis. The spadix is robust and forms a uniform width throughout the 
different zones. The female flowers are green when fresh and this is only 
observed in S. tessellata. 


Etymology: The specific epithet is derived from the name of the type and 
only known locality. 


Studies on Schismatoglottideae 209 


Acknowledgements 


This study is funded by the Ministry of Higher Education, Malaysia by 
fundamental research grant scheme No. FRGS/01(04)/609/2006(42) under 
Sarawak Forestry Department Research Permit No. NPW.907.4.2(I)-101 & 
Park Permit No. 58/20076. The collaboration and support of the Sarawak 
Forestry Corporation, the Forest Research Centre (Kuching), and the 
Sarawak Biodiversity Centre, are gratefully acknowledged. Special thanks 
are due to P. C. Boyce for his comments on the manuscript. 


References 


Hay, A. and Yuzammi. 2000. Schismatoglottideae in Malesia I — 
Schismatoglottis. Telopea 9(1): 1-178. 


Ridley, H.N. 1907. New or rare Malayan plants. III. Journal of the Straits 
Branch of the Royal Asiatic Society 49: 11-52. 


Wong, S.Y. & Boyce, P.C. 2010. Studies on Schismatoglottideae (Araceae) of 
Borneo IX: A new genus, Hestia, and resurrection of Apoballis. Botanical 
Studies 51: 249-255. 


Wong, S.Y., P.C. Boyce, A.S. Othman and C.P. Leaw. 2010. Molecular 
phylogeny of tribe Schismatoglottideae based on two plastid markers 
and recognition of a new tribe, Philonotieae, from the neotropics. Taxon 
59(1): 117-124. 


Gardens Bulletin Singapore 62 (1): 211 -212. 2010. Dati 


Book Review: B.S. Parris, R. Kiew, R.C.K. Chung, L.G. Saw & E. 
Soepadmo (eds). 2010. Flora of Peninsular Malaysia, Series I. Ferns and 
Lycophytes. Vol. 1. (Malayan Forest Records. No. 48). Forest Research 
Institute Malaysia, Ministry of Natural Resources and Environment, 
Malaysia, 249 pp. Price: RM80/USD60 


A new flora useful to the study of Singapore pteridophytes is now available. 
The volume | of the new Flora of Peninsular Malaysia, Series 1, dealing with 
fern and lycophyte groups, has just been published. 

The flora, according to the new publication, was initiated to document 
the indigenous plant diversity by providing reliable and accurate accounts 
of plant families, genera and species found in Peninsular Malaysia, with 
updated nomenclatures. It will consist of two series, with Series IT dealing 
with gymnosperms and angiosperms. 

The Vol. 1 of Series 1 of the new flora described 100 species in 21 
genera and 9 families of ferns and lycophytes, representing about a sixth of 
the spore producing vascular flora of the peninsula. The families treated 
by various authors and included in vol. | are Selaginellaceae (K.M. Wong), 
Psilotaceae (R. Kiew), Equisetaceae (R. Kiew), Osmundaceaee (R. Jaman), 
Matoniaceae (R. Jaman & Y. Umi Kalsom), Schizaeaceae (R. Jaman & Y. 
Umi Kalsom), Cibotiaceae (A.T. Nor Ezzawanis), Loxogrammaceae (R. 
Jaman) and Grammitidaceae (B.S. Parris). All species are shown with a 
distribution map and provided with an assessment of their conservation 
status. But only selected species of each family are nicely and accurately 
illustrated with line drawings. However, each family is further illustrated 
with beautiful coloured photos of a few more representative species. 

There is one nomenclatural novelty proposed in vol. 1 of the new 
flora, namely, Tomophyllum callophyllum (C.H. Wright) Parris, and no 
new species record is reported. The biggest taxonomical change is seen in 
the Family Grammitidaceae where several new generic concepts recently 
proposed based on molecular evidence are accepted. 

Aside from the useful information on the taxonomy and nomenclature, 
the book includes also a conspectus of the orders, families and genera of 
ferns and lycophytes of Peninsular Malaysia, keys to the families, genera 
and species of the plant groups treated, a historical account of botanical 
collecting of ferns and lycophytes in Peninsular Malaysia, and interestingly, 
a chapter on the assessment and conservation of these plant groups in 
Peninsular Malaysia. A glossary and a separate index to the scientific names 
and vernacular/common names of the taxa conclude the volume. 

I have one question in mind after browsing through the hundred pages 
of the flora. Figures A & B in Plate 2 represent two different species? 


PIMP Gard. Bull. Singapore 61 (2) 2010 


As a student of Malesian pteridophyte flora, I definitely look forward 
to the completion of the publication of Series | of this new flora. The authors 
of vol. 1 of Series 1 of the new flora are to be congratulated for this timely 
and well-done accomplishment. 


BC Van 
The Herbarium 
Singapore Botanic Gardens 


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